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BOTANICAL MUSEUM 
LEAFLETS 
HARVARD UNIVERSITY 


PRINTED AND PUBLISHED AT THE 
BOTANICAL MUSEUM 
CAMBRIDGE, MASSACHUSETTS 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


VOLUME XXII 


BOTANICAL MUSEUM 
CAMBRIDGE, MASSACHUSETTS 
1967-1970 


TABLE OF CONTENTS 


NuMBER 1 (September 8, 1967) 


Bat Cave Revisited 
By Pau. C. Manceusporr, Hersertr W. Dick 
AND JULIAN CAMARA-HERNANDEZ ......1 


NUMBER 2 (December 27, 1967) 


Prehistoric Maize, Teosinte, and Tripsacum from 
Tamaulipas, Mexico 
By Paut C. Mancetsporr, RicHarp S. 
MacNeEIsH aND Watton C. GALINAT . . . 33 


NuMBER 38 (November 22, 1968) 


Catalogue of Infrared Spectra of Fossil Resins 
(Ambers) [—North and South America 
By JEAN H. LANGENHEIM AND Curt W. BEcK 65 


Number 4 (January 10, 1969) 


De Plantis Toxicariis e Mundo Novo Tropicale 
Commentationes III. Phytochemical Exami- 
nation of Spruce’s Original Collection of 
Banisteriopsis Caapi 
By Ricnuarp Evans Scuuutes, Bo HOLMSTEDYT 
AND JAN-ERIK LINDGREN. ....... . 121 


De Plantis Toxicariis e Mundo Novo Tropicale 
Commentationes IV 
By Ricuarp Evans SCHULTES. ..... .. . 138 


[v ] 


NumBeEr 5 (June 18, 1969) 


Tree Datura Drugs of the Colombian Sibundoy 
By Mervin L. Brisror . 


NuMBER 6 (June 25, 1969) 

De Plantis Toxicariis e Mundo Novo Tropicale 
Commentationes V. Viro/a as an orally ad- 
ministered hallucinogen 
By Ricuarp Evans SCHULTES . 


Number 7 (November 21, 1969) 

A New Amazonian Arrow Poison: Ocotea venenosa 
By A.J. Kosrermans, Homer V. PINKLEY AND 
Witiiam L. SrerN 


. 165 


. 241 


Blanche Ames Ames (1878-1969)—An Appreciation 


By Ricnarp Evans SCHULTES . 


Four New Species of Saurauia from South America 
By Duasa D. Sorsarro 


Number 8 (December 26, 1969) 
Natural and Artificial Hybrid Generic Names of 


Orchids. Supplement I: 1966-1969 
By Lestie A. Garay AND Herman R. SWEET 


Numser 9 (April 80, 1970) 


Teosinte Introgression in the Maize of the Nobo- 
game Valley 
By H. Garrison WILKES . 


Karly Kight-Rowed Maize from the Middle Rio 
Grande Valley, New Mexico 
By Watton C. Gauinat, THEODORE R. 
REINHART AND THEODORE R. FRIsBIE 


[vi ] 


. 297 


. 318 


NuMBER 10 (June 29, 1970) 


Ethnogynecological Notes in the Harvard Univer- 
sity Herbaria 
By Siri von Reis AttscHuL ..... . . . 888 


De Plantis Toxicariis e Mundo Novo Tropicale 
Commentationes VII. Several Ethnotoxicologi- 
cal Notes from the Colombian Amazon 
By Richarp Evans SCHULTES. ...... . 845 


[ vii | 


INDEX OF ILLUSTRATIONS 


Ambelania Lopezii Woods . . . . UXNNX, LXXX1 
Banisteriopsis Caapi (Spruce ex Griseb.) Morton 
XXIX, XXXI-XXXITI 
Blanche Ames Ames .......... . UXVI 
Blanche Ames Ames and Oakes Ames. . LANNITI 
Calopogon pulchellus (Salisb.) R.Br. 2... . UNIX 
Chelonanthus alatus (lubl.) Pulle . 2 2 1... XL 
Connarus opacus Schellenb,. . 2. 2. 2 2 2. XXXV 
Connarus Schultesii Stand. 2. 2 2... XXAVI 
Cypripedium Calceolus L. 
var. pubescens (Willd.) Correll . . . 2). . UXNXI 


Datura candida ( Pers.) Saff. cultivars 
XLIX-LI, LIT, LV-LXI 


Daturacultivars. .............. Lil 
Datura sanguinea R. & P. cultivars . 2... . LIV 
Davidia involucrata Baill. (Dove Tree) . . LXVII 
Distictella racemosa (Bur. et AK. Schum.) Urb. 

LX XXII 
Duroia hirsuta (P. & #7.) Schum. 2... . . XLII 


[ viii ] 


Duroia petiolaris Hook.f. . 2... . . . . . XXLIV 
Duroia saccifera (Mart.) Benth. & Hook.f. . XLIII 


Kpidendrum tampense Lindl... . . . . . . LXXII 
Evidence of teosinte introgression and maize X 
teosinte hybridization . . ....... LUXXVI 
Habenaria nivea (Nutt.) Spreng. . . . . . . LXX 
Infrared spectraofamber. ..... XIV-NNVIII 
Lisianthus nigrescens Cham. et Schlechtd. . . . XLI 
Maize of the Nobogame Valley ..... . LXNV 


Map of maize-containing sites in New Mexico LXXIX 


Map of the area of the Colombian Amazon 
inhabited by Witoto Indians . . ... . . LXII 


Map of the Valley of Sibundoy, Colombia . XLVIII 


Martinella obovata (H BK.) Bur. et K. Schum. 
LXXXIII 


Ocotea venenosa Kosterm. et Pinkley . . UX1V, LXV 


Polyradicion Lindenii (Lindl.) Garay . . . LXVIII 
Prehistoric maize from Bat Cave .... .. I-VII 


Prehistoric maize from Tamaulipas VIII-NI, XIII 


Prehistoric teosinte from Tamaulipas... . . XII 
Prehistoric Tripsacum from Tamaulipas... XIII 
Psychotria involucrata Sw... .......XLVI 


[ ix ] 


Psychotria nudiceps Stand... 2 2 2... XLV 


Psychotria psychotriaetolia (Seem.) Standl. . XLVII 
Richard Spruce... 2 2. NRX 
Rourea glabra WBA. 2... 2. 2... 6 XXXNVII 
Schoenobiblus peruvianus Stand. . . . . NNXVIITI 
Stelis pendulispica dmes . . . . 2... . LUXNIV 
Styrax Tessmannii Perk. . 2... . 2.) 0 OXNNXIX 
Teosinte-pollinated maize ears . . . . . LXNXVITI 
Tripsacoid characters in maize cobs. . . LXNVIIL 


Unonopsis veneficiorum (Mart.) RB. Fr. . XXXIV 


Virola theiodora (Spr. ev Bth.) Warburg. .  LNITI 


INDEX 
TO GENERA AND SPECIES 


AGATHIS, 79, 83, 89, 90 
australis L., 89 


AGERATUM 


conyzoides, 338 
agua, 204 
agua blanca, 191, 192 
ACTINIDIACEAE, 265 


ALLOPLECTUS 
semicordatus P. & E., 151 


amaraguna, 148 
amaron, 185, 209 


amaron borrachera, 209 


AMBELANIA 
Lopezii Woods. ex R.E. 
Schult., 347-349, 351 

amber, 65-120 


ANADENANTHERA 
colubrina, 237 
peregrina, 230, 237 


andaqui, 185, 223 
andaqui borrachera, 223, 224 
ANNONACEAE, 134, 337 


APHANA 
sp., 337 


APOCYNACEAE, 337, 347 
ARACEAE, 345 
ARAUCARIA, 78, 83 


ARAUCARIACEAR, 78,79,82,83, 89 


ARAUCARIOXYLON, 83 
ARAUCARIACITES 
australis Cook., 78 
ARAUCARITES, 83 
longifolia, 89 
ARISTOLOCHIA 
sp., 341 
ARISTOLOCHIACEAR, 341 
| arrow poisons, 136, 144, 241—- 
251, 347, 351 
ayahuasca, 121, 237 
BANISTERIA, 122 
Caapi Spruce ex Griseb., 122, 
124 
sp., 126, 127 
BANISTERIOPSIS, 122, 158 
Caapi (Spruce ex Griseb.) 
Morton, 122, 123, 126, 128, 
129, 164, 237 
inebrians, 164, 237 
Rusbyana, 164, 230, 237 
spp., 191, 192, 195 
beans, 182, 297 
BESLERIA 
ignea Fritsch., 151 
be-zia, 148 
biangdn, 185, 208 


biangan borrachera, 188, 190, 
208, 209 


biaxit, 195 


[ xi ] 


BIGNONIACEAE, 337 
blanco, 204 
BoraGINAcEAE, 336-338, 340 
borracha, 202 
borracho, 202 
borrachera, 168, 184, 185, 187, 
190, 193, 202, 204, 206 
borrachera de agua, 202 
borrachero, 206, 214 
borracherushe, 202 
BRACHYPHYLLUM- 
PAGIOPHYLLUM, 79 
BRUGMANSIA 
arborea Lagerh., 201 
aurea Lagerh., 201 
bicolor Pers., 198 
candida Pers., 201 
sanguinea D.Don, 198 
bui-ish borrachera, 204, 205, 208 
bui-ish borracherushe, 205 
BULBOSTYLIS 
eapillaris, 339 
BUNCHOSIA, 124 
burundanga, 191 
buyés, 202, 204 
buyés borrachera, 202, 204, 205 
buyés borracherushe, 202 
BYRSONEMA, 124 
caa-pi, 127 
caapt, 121, 122, 124, 126, 127, 
237 
caapi-pinima, 127 
calientes, 190 


CALOPOGON 
pulchellus (Salisb.) R. Br.,259 


CAMPANULACEAE, 341 
CAMPELIA 
zanonia, 338 
CANAVALIA 
sp., 341 
cari, 185, 214 
cari borrachero, 213 


CENTROPOGON 

calycinus, 341 
CEPHALOTAXUS, 73 
chalua borrachero, 214 
CHELONANTHUS, 151 

alatus (dubl.) Pulle, 146, 147 

chelonoides (L.) Gilg., 148 
chicha, 193, 347 
chontaruco, 185, 208 
chontaruco borrachera, 208 
CoMMELINACEAR, 338 
ComposiTak, 336, 338, 339 
CoNNARACEKAR, 136, 138 
CONNARUS 

lentiginosus Brandg., 140 

opacus Schellenb., 136, 137 


Schultesii Standl. ex Schult. , 
138-140 


Sprucei Baker, 137 
CONOMORPHA 
citrifolia Mes, 346 


CREMASTUS 


QQ" 


sceptrum, 33; 


CRYPTOCARYA 
Bowiei (Hook. ) Druce, 244, 
246 


cuauhchichic, 134 


cucu, 185, 211 


[ xii | 


culebra, 185, 220 


culebra borrachera, 185, 189- 
191, 220, 222 


cumala, 232 

CUNNINGHAMITES, 83 

Cupressackeak, 79, 82 

CUPRESSINOXYLON, 82 
Bibbinsii, 82 

curare, 136, 144, 351 


CYPERUS 
brevifolius, 341 


CYPRIPEDIUM 
Calceolus L. 
var. pubescens (Willd. ) 
Correll, 261 


CYRTANDRA 
Cumingii, 337 
sp., 337 
DAMMARA, 83 
australis Lamb., 89 
microlepis Heer, 80 


danta, 185, 208 


danta borrachera, 190, 208 


DATURA, 160, 168-170, 176- 
181, 184, 187-193, 195-198, 
205, 212, 213, 215, 218, 223 


affinis Saff., 201 
arborea R. & P., 201 
aurea Lagerh., 201 


c.v. Andres’, 174, 179, 183, 
186, 189, 198, 218, 222,223 

candida (Pers. ) Saff.,170,172, 
173,175,176, 178, 180-182, 
184,190,197,200-203,218, 
222,223 


[ xiii | 


c.v. Amarén’, 172-176, 178- 
180,183,186,188,198, 209, 
210 

e.v. Biangan’, 172, 174-177, 
179,183,186, 187,198, 206— 
208 

c.v. Buyés’, 168-180, 182, 
183, 186, 187, 198, 202, 
203, 205, 209 

c.v. Culebra’, 170, 172-176, 
179, 183, 186-190, 192, 
193,195,198,205,218-221 

c.v. Dientes’, 173-177, 179, 
180, 183, 186, 187, 198, 204 

e.v.Munchira’,172,174,175, 
179, 183, 184, 186, 189, 
198-195, 198, 213,216,217 

e.v. Ocre’, 172-175, 179, 183, 
186,187,198, 206 

e.v. Quinde’, 172, 174-181, 
183, 188, 189, 193, 198, 
213,215 

c.v. Salaman’, 174, 175, 179, 
183, 184, 186, 188, 193, 
198, 212, 2138 

cornigera, 196 

dolichocarpa (Lagerh.) Saff., 
223 

Pittieri Saff., 201 

sanguinea R. & P., 168, 170, 
196, 198, 204, 220 
c.v. Guamuco’, 179, 183, 

186, 187, 197, 199, 200 
c.v. Sangre, 179, 183, 186, 
187, 197, 201 

sect. Brugmansia, 166 

Stramonium, 181, 218 

suaveolens H, & B. ex Willd., 
208, 218, 220, 228 

vuleanicola 4.8. Barkley, 170 


DAVIDIA 


involucrata Baill., 257 


DAVILLA 
lacunosa, 341 
DEHAASIA 
triandra, 251 
DESMOS 
Hancei, 337 
DILLENIACEAR, 341 
Dirrrera, 99 


DISTICTELLA 
racemosa (Bur. & K. Schum. 
ex Mart.) Urban, 347, 350, 
351 
DUBOISIA, 196 
DUROIA, 151 
hirsuta (P. & E.) K. Schum., 
152-154 
kotchubaeoides Steyerm., 
154, 155 


petiolaris (Spr.) Hook f., 154, 


-_~ 


saccifera (Mart.) Hook,f., 
155, 156 

Sprucei Rusby, 156 
DYALANTHERA 

parviflora, 233 
ELEUTHERINE 

bulbosa, 33 

plicata, 339 
epend, 237 
EPIDENDRUM 

tampense Lindl., 262 
EPIGYNUM 

Maingayi, 337 
ethnogynecological plants, 333- 

343 


EUPHORBIACEAE, 142, 336, 340, 
341 


FARADAYA 
sp., 337 


fish poisons, 1388, 140, 144, 347 
FLACOURTIACEAE, 142 
floripondio, 185, 204 
floripondio blanco, 202 
Jloripundo, 202 

FRANSERIA 


ambrosioides, 339 


GARRYA, 134 
laurifolia (Hartw.) Benth. , 134 
var. macrophylla (Hartw. ) 
Wangerin, 134 


GARRYACEAR, 134 


GEINITZIA, 83 
Reichenbachii (Geinits) Holl. 
& Jeff., 82 


GENTIANACEAR, 146 
GesNERIACEAL, 151, 337 
gingive-k’ 0, 2438 
GLOCHIDION 
cauliflorum, 340 
GLYPTOSTROBUS, 94 
GRAMINEAE, 336, 339, 341 
guambia, 200 
guamuco, 185, 200, 201, 204 
guamuco blanco, 185, 202 
guamuco borrachera, 200, 201 


» 


guamuco floripundo, 185, 202 


GUSTAVIA 
Poeppigiana Berg. ex Mart., 
233 
HABENARIA 


nivea (Nutt.) Spreng., 260 


[ xiv | 


HAEMADICTYON, 127 
hallucinogens, 121-132, 164, 
168, 188-192, 229-240 
HAPLOPAPPUS 
spinulosus, 339 
HELIOTROPIUM 
argenteum, 340 
he-rog, 233 
HINTONIA 
latiflora, 338 
ho-ko-so-gi-nd, 146 
huilca, 237 


HYMENAEA, 92-94, 96, 99, 
100 


Courbaril L., 92-94, 96, 99, 
100 


HYOSCYAMUS, 196 

HYPODAPHNIS 
Zenkeri, 251 

IcAcINACEAE, 337 


INAPERTUROPOLLENITES 
dubius (Potonie & Venitz) 
Thompson & Pflug, 79 
inzo?tsi, 246 
IOSTEPHANE 
heterophylla, 339 


IRESINE 
celosia L., 184 
herbstii Hook,f., 184 


[RIDACEAE, 339 
i-te-si-fan-di, 136 
JATROPHA 
angusti, 341 
JUNIPERUS, 73 
hynoides Heer, 80 


KADSURA 


scandens, 336 
kinde-borrachera, 220 


KNEMA 


glomerata, 337 
kuku, 211 
kurru, 234 
kutrucu, 234, 236 
LasraTak, 336, 340 
LARIX, 73 
LauRAcEAR, 241, 244, 250, 251 
LAURUS 
nobilis, 251 
lee-the, 151 
LrauMInosak, 92, 336, 340, 341 
lengua de tigre, 185, 190 
le-sa, 233 
Litiaceak, 336 
LINDERA 


Benzoin, 251 
LIQUIDAMBAR, 73, 86 
LISIANTHUS 

nigrescens Cham. & Schlecht., 

146, 148, 149, 151 

pendulus Mart., 151 


LoGANIACEAER, 339 


LorANTHACEAE, 340, 341 


LORANTHUS 
sp., 340, 341 


lulumoco, 267 


LUNANIA 
parviflora Spruce ex Benth. ,142 


maicillo, 298 


[ xv ] 


maize, 1, 2, 6, 33, 34, 36, 40, 
42-44, 47-49, 54, 57, 58, 62, 
182, 297-301, 303-3807, 309, 
310, 313 
archeological, 313-329 
prehistoric, 1-24, 33-49, 52- 

63, 310 

maisillo, 297 

matsmillo, 297 

mais silvestre, 298 

maleficio, 190 

MALLOTUS 
Poilanei, 341 

MALPIGHIACRAE, 122, 124, 127, 
338 

MARTINELLA 
obovata (HBK.) Bur. et K. 

Schum, ex Mart., 347, 351, 
352 

MASCAGNIA 
septentrionalis, 338 

mata cachorro, 140 


mata negro, 140 


matamata, 233 


MAURITIELLA 
aculeata (HIBK.) Burret, 236 
medicinal plants, 134,138,140, 
142, 146, 148, 151, 168, 184, 
186, 188, 189, 197, 211, 333- 
351 
mee-tsee-boo-koo'na, 347 
MELICOPE 
monophylla, 337 
me-re-ta-kee, 144 
METAPLEFENTICIRAS 
pacificum Smith, 90 


METASEQUOIA, 94 


METHYSTICODENDRON 
Amesianum R.E. Schult., 195, 


218, 222 


MIMOSA 
hostilis, 230, 237 


molongo, 347 
mongajo, 208 
mtzkway borrachera, 222 
munchira, 185, 214, 216 
munchira borrachera, 216 
munchiradas, 216 
mutscuat, 185,218 
mutscuat borrachera, 220 
MyrisTicackak, 387 
MyrsINnaceaAk, 346 
NAUTILOCALYX 
sp., 151 
NECTANDRA 


coriacea, 251 

globosa, 251 

Rodiet, Rob. Schomb., 243 
ngunsiana, 185, 214 
ngunsiana borrachera, 2138 
NEOLITSEA 

Levinei, 251 
NEONAUCLEA 

formicaria, 337 
NEPETA 

Cataria, 340 
nyakwana, 230, 237 


OCIMUM 


sanctum, 340 


[ xvi | 


OCOTEA, 243, 251 
palmana, 251 
Rodiei (Rob. Schumb.) Mez, 

243, 244 
venenosa Kosterm. & Pinkley, 
241, 244-246, 250, 251 

OvacackEAk, 341 

6-me’-na, 346 

00-kod-he, 234 

oo-kob-na, 232, 233 

o-pri-lo, 164 

orchid hybrids, generic names 
of, 273-289; parentage of, 
289-296 

o-sia, 148 

PALICOUREA, 164 

parica, 237 

PAULLINIA, 124 

payé, 234 

PENNISETUM 


alopecuroides, 341 


PHASEOLUS 
adenanthus, 340 


PHENAX 


integrifolius Wedd., 187 


PHYLLANTHUS 
lathyroides HBK., 142 


PHYTOCRENE 


Blancoi, 337 
PICEA, 73, 83 
PrnacEak, 73, 75, 79, 80, 83 
pinolilos, 148 


PINUS, 78, 82, 83 
sp., 80 


PIpeRAckAk, 338 
PITYOXYLON, 80, 83 
PopocarRPackEak, 79, 82 


poisonous plants, 142, 144,151, 
152, 154, 156, 158, 184, 346 
POLLIA 
thyrsifolia, 338 
POLYOSMA 
sp., 337 
POLYPREMUM 
procumbens, 339 


POLYRADICION 
Lindenii (Lindl.) Garay, 258 


POTOMORPHE 
peltata, 338 


PREMNA 
sp., 341 

PREPINUS, 82 

PROTIUM, 98 
Icicariba, 98 

PSYCHOTRIA, 156, 164 
carthaginensis Jacq., 158,159 
involucrata Sw., 158, 161 
nudiceps Standl., 158, 159 
psychotriaefolia (Seem. ) 


Standl., 158,163, 164,230, 
237 


psychotropic plants, 168, 185, 
188-197 


PTEROCARPUS 
indicus, 340 


PTYCHOPETALUM 


olacoides, 341 
quinde, 185, 214 
quinde borrachera, 213, 214 


RAMONDIA 
pyreaica Rich., 151 


[ xvil | 


RAVENSARA Reichenbachii (Genitz) Heer, 


crassifolia, 251 80, 82 
resfrio, 189 SEQUOIADENDRON, 73 
ROUREA sha-ka-ker-nda-se, 152 
erecta (Blanco) Merr., 140 hiaainde, 142 


glabra HBK., 140, 141 i 
ligulata Bak., 142 mlenBeiee- 1805, Tp 


: , SoLANACERAR, 220 
Rupraceak, 151, 336-358 , 


soliman, 152, 154 
SPIGELIA 
pedunculata HBK., 200 


Ruracrear, 337 


SABICEA 
Vogelii, 338 

squash, 297 

STELIS 


pendulispica Ames, 264 


salaman, 185,212 
salaman borrachera, 212, 213 


salamanga, 185 Buc caeacie. Lal 
: ACACKAE, 


salamanga borrachera, 212 ee : 

eee ear , STYRAX 

salvanje, 185 Tessmannii Perk., 144, 145 
salvanje borrachera, 212 TAXAcEAE, 73 

SALVIA TaxopiackAk, 73, 75,79, 80, 82, 


83, 94 
TAXODIUM, 73 


teosinte, 1, 4,9, 12, 16, 34, 37, 
40, 45-49, 61, 62, 297-301, 
303-307, 309, 310, 319 


THEOBROMA, 236 


divinorum, 340 
SAPINDACEAE, 124 
SAURAUIA, 265 
chaparensis Soejarto, 270 
Mexiae Aillip ex Soejarto, 268 


multinervis Soejarlo, 266 


Schultesiana Soejarto, 267 'THYMELEACEAR, 142 
SAXIFRAGACEAR, 337 tinye, 190 
ScHISANDRACEAR, 336 | TORREYA, 73 
SCHIZOSTACHYUM TOURNEFORTIA 


Lumampao, 336 brevilobata, 337 


volubilis, 338 
SCHOENOBIBLUS 
| trago, 193 


TRIPSACUM, 12, 16, 34, 37, 


peruvianus Standl., 142-144 


SEQUOIA, 73, 80, 82-84 47, 48, 62 
ambigua Heer, 87 dactyloides, 48 
heterophyllus, 80 | zopilotense, 48 


[ xviii | 


tsushie borrachera, 206 peruviana (4.DC.) Warb. ,236 
theiodora (Spr. ex Benth.) 


ucuna, 234 
Warb., 229, 230, 232, 235, 


una de tigre, 148 237 
veneficiorum ( Mart.) R. E.Fr., lomatophylla, 339 
134-136 
UROSPATHA, 345 VocHYSIACEAE, 339 
caudata Schott, 346 WIDDRINGTONIA 
sagittaefolia Schoti, 346 Reichii (EH.) Heer, 80 


somnolenta I.E. Schult., 346 
wy-gaw-o-meé-ki, 347 
VERBENACEAR, 337, 341 
wy-soo-dé, 158 


VIGUIERA 
montana, 339 yagé, 191 
vinho de yurema, 237 yaje, 164, 237 
VIROLA, 229, 230, 232-234, yakee, 237 
236, 237 P 
calophylla Warb., 229, 230 Ger Aa 
calophylloidea Markg., 229, yoom-dd-ka, 347 
230, 236 _ 
elongata ( Benth.) Warb., 229, al 
236 yurema, 237 


[ xix ] 


ERRATA 


Page 336, line 6 
for twelve read fourteen 


Page 336, line 8 
delete ‘‘the Gramineae by four,”’ 


Page 339, line 6 
for Gramineae read Cyperaceae 


Page 340, last line 
replace the , by a. 


Page 341, line 1 
delete ‘‘from the Gramineae’’ 


Page 341, line 2 
insert , Gramineae after Frake 512 


Page 341, line 4 


insert , Gramineae after Kajewski 276 


Issued January 26, 1972 


[ xx ] 


BOTANICAL MUSEUM LEAFLETS 


HARVARD UNIVERSITY 


CampripGr, Massacuusetrts, SEPTEMBER 8, 1967 Voi. 22, No. 1 


BAT CAVE REVISITED 
BY 
Paut C. Mancetsporr, HERBERT W. Dick 
AND JULIAN CAMARA-HERNANDEZ* 


In 1948 an expedition sponsored by the Peabody Mu- 
seum of Harvard University and led by Mr. Herbert 
Dick, then a graduate student, uncovered many cobs and 
other parts of maize from the accumulated refuse in a 
once-inhabited rock shelter in New Mexico known as 
Bat Cave. These prehistoric remains were significant in 
several respects: (1) they were the first remains dis- 
covered which showed clearly the nature of primitive 
maize; (2) they provided tangible evidence of a well 
defined evolutionary sequence; (3) they represented the 
first archaeological evidence showing: (A) that primitive 
maize was both a popcorn and a form of pod corn, (B) 
that maize did not originate from teosinte, (C) that much 
of the variation in modern maize is the product of intro- 
gression from teosinte (Mangelsdorf and Smith, 1949). 

Since the 1948 expedition had excavated only one sec- 
tion of the eave, Mr. Dick in 1950 made a second ex- 
pedition which excavated additional sections in the hope 
of finding still older vegetal remains. A_ preliminary 


* The authors are respectively Fisher Professor of Natural History, 
Harvard University; Associate Professor of Anthropology, Adams 
State College, Alamosa, Colorado; and Fellow of the Consejo Nacional 
de Investigaciones Cientificas y Técnicas de la Argentina. 


[1] 


examination of the maize specimens of this second ex- 
pedition showed that the earliest cobs were even more 
primitive in some respects than those found in the 1948 
excavation and the specimens tended to confirm in gen- 
eral the principal conclusions drawn from the detailed 
study which had been made of the 1948 collection. It 
was our intention at that time to make a similar detailed 
study of the 1950 collection and to publish the results at 
an early date. In the meantime, however, the senior 
author participated in a project involving the classifica- 
tion and description of the living races of maize of Mexico 
(Wellhausen et al., 1952) and this made it possible to 
identify precursors of several living races of maize found 
in archaeological excavations in northeastern Mexico by 
MacNeish (Mangelsdorf et al., 1956) and in northwestern 
Mexico by Lister (Mangelsdorfand Lister, 1956). Studies 
of still other collections of archaeological maize had shown 
strikingly the importance of teosinte introgression in the 
evolution of maize (Galinat et al., 1956). In view of 
these developments, it was decided to postpone the final 
studies of the 1950 collection of Bat Cave specimens until 
still other collections which had come to the Botanical 
Museum from various sources could be analyzed. These 
several studies of specimens from sites in Mexico and the 
United States have been completed (Galinat and Ruppe, 
1961; Galinat and Gunnerson, 1963; Mangelsdorf et a., 
1964; Mangelsdorf et a/., in press). We can now re- 
examine the two Bat Cave collections with the benefit 
of a familiarity with the living races of maize, a con- 
siderably greater experience in analyzing archaeological 
material, and in the light of significant evidence revealed 
by other collections of which one of the most important 
is from sites in the Valley of Tehuacin in Mexico in 
which remains of prehistoric wild corn were found. 


[2] 


THe 1950 ExcavATION 


Since the Bat Cave site and its environs were described 
in detail in the 1949 report as well as in a more recent 
publication (Dick, 1965) there is no need to repeat the 
description here. 

The 1950 expedition comprised three members all from 
the University of Colorado; Herbert W. Dick, who 
directed the excavation, Francis Olson and Allen Olson. 
The excavation was made from August 1 to September 6. 

The main purpose of the second expedition was to 
obtain additional prehistoric vegetal material, especially 
of maize, and charcoal from all levels for radiocarbon 
dating. Stone and other artifacts were collected as by- 
products. Except for finding a new type of knotted-fiber 
container these proved to be similar in virtually all re- 
spects to those turned up in the 1948 excavation. 

Because there was no obvious stratigraphy in the cave 
corresponding to cultural phases, the material was re- 
moved in arbitrary strata of 12 inches each: 0-12’, 12- 
24/’, ete. Since three different sections of the cave were 
excavated and since the sterile sand representing Antevs’ 
dry level on which the cave deposits rest, occurs at dif- 
ferent levels in different parts of the cave, it might be 
supposed that arbitrary levels in one section would not 
correspond to those in another. In Section IB for ex- 
ample, the sterile sand occurs at 86’’ while in IC it is 
found at 60’. However, the cobs in the 24-36” level of 
IB are similar to those in the 24-86” level of other sec- 
tions. Consequently we have averaged the data on cobs 
and kernels according to the arbitrary levels in which the 
specimens were found. Although this procedure lacks 
refinement it still demonstrates an evolutionary sequence 
of the maize remains from the lower to the upper levels 
of the deposit. 


[ 3 ] 


Dating of the Remains 

One of the most important questions to be answered 
is the date of the remains. The radiocarbon determina- 
tions (Libby, 1951; Arnold and Libby, 1950) of samples 
of charcoal and other materials from the several levels 
are set forthin Table I. The data from the two excava- 
tions are fairly consistent for the three uppermost levels 
but differ widely for the two lower levels. There is reason 
to believe that the date of 5605 for the charcoal in the 
48-60” level is not valid for the associated earliest maize. 


Tas_e I. Radiocarbon dates of charcoal, wood fragments, and cobs 
from the two Bat Cave excavations. 


1948 Excavations 1950 Exeavations 
Material Level Date Material Level Date 

cobs 0-12” |1752+250 

wood 12-24" | 1907+250] Charcoal 11-15” | 1610+200 
wood 24-36" | 2239+ 250 ” 24-86" | 2816+ 200 
cobs, wood | 36-48/" | 2249+250 “ 36-48” | 20484170 
wood 48-60” | 2862+250 “ 48-60” | 56054290 
Peres ies 60-66" | 3000-3500 = 60-66” | 593814310 
ation 


On the basis of the characteristics of the cobs, especially 
those which represent evidence of teosinte contamina- 
tion, we have concluded that the maize from the 48-60” 
level of Bat Cave is later than the maize of the Abejas 
phase in the Tehuacan caves and earlier than that of the 
Ajalpan phase. This would date it at between 2300 and 
1500 B.C. A similar correlation with the remains of 
maize in Romero’s Cave, Tamaulipas, Mexico, excavated 
by Dr. Richard S. MacNeish (in press) makes it con- 
temporary with the Guerrero phase dated by radiocarbon 
at 1800-1400 B.C. These correlations suggest that the 
earliest Bat Cave maize is probably not earlier than 


[ 4 ] 


ca 2300 B.C. and may be several centuries later. 

The 1950 Bat Cave collection comprises 816 specimens 
and includes cobs, eroded fragments of cobs, kernels, 
husk systems and husk fragments, peduncles, pieces of 
stalk, and one tassel fragment. We shall consider these 
below in relation to the levels in which they were found 
in the Cave. 

Tue Cops 

On the whole the cobs from this collection, numbering 
444 specimens in addition to one fragment bearing ker- 
nels, were quite similar to those of the earlier excavation 
but with one important difference. Some of the cobs from 
the lowest levels were smaller and more primitive than 
any of those previously turned up at this site. Three of 
these are illustrated in Plate ILI, A. 


Early Maize a Popcorn 


The senior author and his associate, Dr. Walton C. 
Galinat, made an intensive study of one of these speci- 
mens which contained the partial remains of a single 
kernel. We spent several days in dissecting this cob and 
measuring all of its parts. On the basis of these measure- 
ments, Galinat prepared the diagrammatical longitudi- 
nal section illustrated in Fig. 1. We concluded that the 
tiny kernels which this cob must once have borne could 
only have been those of popcorn, a type in which the 
kernels are small and hard and are capable of exploding 
when exposed to heat. This conclusion has been amply 
confirmed by finding among the prehistoric grains several 
actual specimens of popped corn described later. The 
Bat Cave specimens thus provide convincing archaeolo- 
gical evidence in support of one part of the conclusion 
first reached by Sturtevant (1894) and later by Mangels- 
dorf and Reeves (1939) that primitive corn was both a 
popcorn and a pod corn. 


[5] 


Early Maize a Pod Corn? 

Whether the earliest Bat Cave corn was also a pod 
corn depends upon how pod corn is defined. Geneticists 
know that the principal characteristics of pod corn are 
controlled by a single locus, 7'u, on chromosome 4. 
Mangelsdorf and Reeves (1989) considered the Tu gene 
to be an ancient wild one still surviving in modern varie- 
ties but they also pointed out that the pod corn of today 
is not the pod corn of wild corn: 


The pod corn found occasionally as a mixture in modern culti- 
vated varieties iscertainly not the wild pod corn which the wander- 
ing Indians discovered millenia ago. Modern pod corn is the result 
or superimposing a single ‘wild’ gene with perhaps a few closely 
linked modifiers upon a genetic complex which has been tre- 
mendously altered by centuries of domestication. 


Intensive research on pod corn and its genetic locus 
by the senior author and his associates over a period of 
almost 80 years has shown that 7'w locus is a compound 


j 


» 


—€ 


re : Mi 
F a | 


Fic. 1. Diagrammatic longitudinal section of one of the Bat Cave cobs based 
on measurements of dissected parts. The tiny kernels show that this was a pop- 
corn; the long pedicels on which the kernels are borne and the bracts which 


almost enclose them indicate that it was also a pod corn. 
Drawn by W.C. Gatinat 
[6 ] 


one having at least two components (Mangelsdorf and 
Galinat, 1964) and possibly three, one of which is iden- 
tical in its expression with a previously recognized mu- 
tation at this locus designated as ‘‘half-tunicate’’ because 
its effects are about half of those of tunicate (cf. Mangels- 
dorf and Reeves, 1959). We also discovered in the pop- 
corn variety, Baby Golden, a major modifying gene 
which we have called tunicate inhibitor and given the 
symbol, 7%. This gene has its locus on chromosome 6 
and is linked with the gene for endosperm color on that 
chromosome. Combinations of the several components 
of the 7'u-tu locus with and without the major modifying 
factor 7% produce a spectrum of phenotypes at one end 
of which all of the kernels are completely enclosed in 
glumes and at the other all of the kernels are completely 
exposed in their upper surfaces but are surrounded in 
their lower surfaces by glumes which are longer than 
those of modern varieties of corn. 

The earliest cobs from the Bat Cave 1950 collection 
are of this latter phenotype as the diagram in Fig. 1 
shows. They have relatively long floral bracts, the lem- 
mas and paleas surrounding the kernels but net com- 
pletely enclosing them. In the 1950 collection we have 
also found one fragment of a cob containing kernels 
which shows exactly this condition. This is illustrated in 
an enlarged photograph in Plate II, C, D, and in Fig. 2. 

The earliest cobs also have other characteristics of pod 
corn including the slender central stem, the rachis, and 
relatively long secondary stems, the pedicels or rachillae, 
upon which the kernels are borne. 

There has been some skepticism with respect to our 
conclusion that cobs of this type represent pod corn 
(Randolph, 1956; Weatherwax, 1956; Goodman, 1965). 
Since prehistoric kernels from such cobs, even when 
present, have long since lost their viability there is no 


[7] 


way of determining their genotype, However, since we 
can duplicate the principal characteristics of such speci- 
mens by combining the components of the Zu locus on 
chromosome 4 with the major modifying factor 7% on 
chromosome 6 and with other modifying genes from 
various popcorn varieties and since neither we nor our 
critics have yet found any other way to duplicate these 
characteristics, we shall continue to regard these early 


Fic. 2. Diagrammatic cross section of the early Bat Cave cob 


od 
KA FT illustrated in Plate II, C and D. The slender rachis (inner 
ORNS No circle), the long rachillae (distance between rachis and base of 
, a ie Ly kernels), the long glumes (solid line), and the paleas (broken 
" . /--\ NY ) line) which almost enclose the kernels, all indicate that this is 
SE a form of pod corn. 


cobs as representing a form of pod corn and we shall ad- 
here to the conclusions of Sturtevant and of Mangelsdorf 
and Reeves that primitive corn was both a popcorn and 
a pod corn. 


Changes in the Rachis Diameter 


We shall, however, modify the conclusion reached in 
the description of the cobs from the 1948 collection in 
which we attributed changes in the cob/rachis ratio 
largely to replacement of higher alleles of the T'u-tu 
locus by lower alleles including one designated as tu”. 
We have not in our experimental cultures been able 
definitely to establish the existence of such an allele or 
if it exists to distinguish its effects from those of various 
modifying factors whose role, in causing variation in the 
diameter of the rachis and the length of the glumes, was 
postulated in the 1949 paper. Changes in the cob/rachis 
ratio in this material now appear to be more a matter 
of increase in the diameter of the rachis than in the re- 
duction in the length of the glumes. The data in Table 
II, based on 430 cobs in which the rachis diameter could 
be measured, show that there has been an increase in the 


[8] 


rachis diameter from 7.0 mm. to 8.9 mm. in the period 
represented by the remains of maize in this cave. 


Increase in Variation 


More conspicuous than the increase in the size of the 
rachis is the increase in variation of the cobs. Several of 
the earliest cobs from the 1950 collection resemble the 
prehistoric wild corn from the Tehuacdn site but not all 


Tasie II. Rachis diameters (mm.) and kernel-row-numbers of the 
cobs from successive levels of Bat Cave. 


Level | Rachis Kernel-row-numbers 
Inches | Diam.Av.|No.Cobs| —g 10 12 14 16 18 | Av. 
0-12 8.9 70 8 20 28 12 2 11.4 
12-24 7.8 99 10 25 35 22 4 3 {11.8 
24-36 7.9 120 12 30 55 16 2 {10.2 
36-48 7.2 142 28 42 54 13 5 10.9 
48-60 7.0 13 | 9 1 g 10.6 
No.Cobs) 430 | 444 | 59 |126 [173 | 65 | 16 | 5 | 


of them are of this type. The majority of them are small 
but there is considerable variation in their other charac- 
teristics, especially in the size and shape of cupules. 
There is some evidence that introgression with teosinte 
had already occurred in some of the cobs from the lowest 
level. Based on the characteristics of the cobs we con- 
clude that the corn from the 48-60” level in Bat Cave, 
as already mentioned, is later than the corn of the Abejas 
phase in the Tehuacan sites and earlier than that of the 
Ajalpan phase. 

The evidence of teosinte contamination is even more 
clear-cut in the specimens from the 86-48” level than in 
the 48-60” level. Photographs of several specimens with 
indurated tissues and solitary pistillate spikelets are illus- 
trated in Plate I, C and E. 

The cobs in this and subsequent levels are so hetero- 


[9] 


geneous in their characteristics that it is impossible to 
classify all of them with respect to race. However, some 
of the cobs clearly resemble those of the race Chapalote. 
A series of these from the lower to the higher levels is 
shown in Plate III, B. 

Another type of cob thicker and more nearly cylin- 
drical in shape than those resembling Chapalote is illus- 
trated in Plate LV. These may be related to the Mexican 
race, Blando de Sonora, described by Wellhausen et a/., 
1952. Some of the kernels with colorless pericarp and 
floury endosperm illustrated in Plate III, C may also be 
related to this race. 

Changes in Kernel-row Number 

An increase in the diameter of the rachis provides a 
larger surface on which kernels can be borne and this 
results in either a higher kernel-row number or in larger 
kernels or in both. ‘Table LI shows that there has been 
a change in the kernel-row number from the lower levels 
to the higher. ‘The modal number in the lowest level is 
10 and in all higher levels, 12. The average row number 
of the cobs from successive levels, the lowest to the 
upper, is 10.6, 10.9, 10.2, 11.8, and 11.4. The increase 
in kernel-row number has, however, been relatively 
smaller than the increase in kernel size discussed below. 


Tue KERNELS 

Increase in Size 

There are 299 well preserved kernels in this collection, 
a much larger number than found in the 1948 material. 
The data in Table III show that there is a substantial 
increase in kernel length and width from the lower levels 
to the higher accompanied, however, by a decrease in 
thickness. The kernels from the lowest level are almost 
isodiametrical in their dimensions; kernels from upper 
levels tend to be longer, wider, and flatter. 


[ 10 ] 


as, | 
a 
a_i 
(eae 


Tare III. Dimensions of kernels of different pericarp and aleurone colors from successive levels of Bat Cave. 


Cave 
Level 


Inches 


0-12 
12-24 
24-36 
86-48 


48-60 


Number of Kernels and their Length, Width and Thickness in mm. 


Brown Red Variegated Colorless Aleurone Color | Averages 
No. L W T |INo. L W TINo. L W TINo. L W TINo. LW T |No. L WT 
7 7.5 7.8 5.5] 1 7.4 11.3 6.9 10 8.2 8.6 5.4 | 18 7.9 8.2 5.5 
446.9 6.8 5.4/10 7.6 7.1 5.3 | 4 8.0 7.8 5.3 | 74 7.8 8.1 5.6 2 8.4 8.5 6.0 |134 7.5 7.6 5.5 
16 8.7 6.7 9:5 S30: 7:8 3.6 19 6.9 6.9 5.5 
29*6.9 6.7 5.7 |26 7.8 6.8 5.3| 9 7.2 7.3 5.5 37 7.1 6.7 5.5 101 7.1 6.8 5.6 
10 6.8 6.7 5.8} 17.7 6.5 5.8 | 11 6.9 6.7 5.8 


* Ten additional brown kernels employed in a popping experiment are not included in these measurements. 


Changes in Pericarp Color 


The majority of the kernels are so well preserved that 
it is possible to classify them with respect to pericarp 
color. Of the 2938 kernels so classified: 116 are brown, 
38 red, 138 variegated, 2 with colored aleurone, and 124 
with colorless pericarp and aleurone. 

The kernels with brown pericarp are more frequent in 
the lower levels than in the higher and they are smaller 
on the average than the kernels with red or colorless 
pericarp. The fact that the kernels from the lower levels 
of Bat Cave are predominantly brown suggests that this 
is the primitive or “‘wild’”’ color. Also most of the early 
prehistoric corn in South America has brown pericarp 
color as do the kernels of corn’s relative, teosinte and 
Tripsacum. 

Pericarp color in maize is the product of the inter- 
action of genes at two, in some cases three, different loci. 
Brown pericarp results when the P factor on chromosome 
1 interacts with one of the several alleles at the A locus 
on chromosome 8: A”, a”, anda, Since the kernels 
from the lower levels are predominantly brown and since 
the a, allele is not common in maize, it seems probable 
that the genotype of the early kernels is 4" 4%PP. The 
red kernels probably represent mutations to 4, perhaps 
resulting from the introgression of teosinte which is 
known to have mutagenic effects (Mangelsdorf, 1958). 

Variegated Pericarp. Thirteen of the 299 kernels had 
variegated pericarp. This variegation is the product of 
a mutation system which has been intensively studied 
by Brink and his associates and which involves the inter- 
action of genes at the P locus on chromosome 1 with a 
genetic factor designated as ‘‘modulator’’ which may 
occupy various positions on chromosome | or on other 
chromosomes. The fact that nine of the 13 variegated 


[ 12 ] 


kernels occur in the 86-48” level indicates that a muta- 
tion system similar to that studied by Brink was already 
in existence at an early date and the fact that the varie- 
gated kernels are larger than either the brown or red 
kernels suggests that the modulator system may have 
caused or been accompanied by other mutations involv- 
ing increases in size. 

Colorless Pericarp. The kernels with colorless pericarp 
are generally larger than those with colored pericarp and 
are predominantly floury. They may belong to the race, 
Blando de Sonora, described by Wellhausen et al., 1952. 
We have already suggested that certain cobs may belong 
to this race. 


Popped Kernels 


In addition to the 293 kernels classified with respect 
to color there were six popped or partially popped ker- 
nels in which the color could not be determined with 
certainty but which appeared to have brown pericarp. 
One of these is illustrated in Plate III, D. We were able 
by a simple experiment to demonstrate that the early 
Bat Cave kernels were capable of popping. ‘To increase 
their moisture content, we placed ten of the smaller 
brown kernels from the 86-48” level in a petri dish with 
a piece of moist paper towel for 48 hours. When dropped 
into hot oil all of these popped in varying degrees. One 
of these, a popped-prehistoric kernel, is illustrated in 
Plate III, D where it is compared with a prehistoric- 
popped kernel from the same level. 


PEDUNCLES 


Twenty-one specimens of peduncles or ‘‘shanks’’ were 
found in this collection. Those from the lower levels 
were usually more slender than those from the higher 
levels as is shown by the specimens illustrated in Plate 


[ 13 ] 


V. Part of the increase in size may be the result of clus- 
ters of small ears being replaced by single large ears as 
discussed below. 
Husks 

The most important modification of a conclusion based 
on the 1948 collections, which resulted from a study of 
the 1950 specimens, is concerned with the nature of the 
husks. There were three husk systems and 47 husk frag- 
ments. In the 1948 collection we found in Stratum II 
a long husk which shows no evidence of ever having been 
shaped around an ear. Furthermore its peduncle as well 
as the peduncle of the ear which it once bore are both 
quite slender. A cob which might have been enclosed in 
this husk system was illustrated in Plate X XVII of the 
1949 report. Because this cob is much shorter than the 
husks, we reached what we then considered an ‘‘almost 
inescapable conclusion”’’ that husks at this stage in the 
evolution of maize did not tightly enclose the ear but 
instead were involucres of leaf sheaths, not greatly modi- 
fied, surrounding the base of the pistillate inflorescence 
but not completely enclosing it. ‘‘Inescapable conclu- 
sions’’ sometimes turn out, when additional evidence is 
forthcoming, to require modification. The additional 
evidence from the 1950 collections suggests that a long 
husk of this type enclosed not a single ear but a cluster 
of ears. ‘This evidence comes from two specimens: one 
husk system (Plate V, B) in which the husks are only 
slightly longer than the 1949 cob which is illustrated in 
Plate VI, Bb and the other a piece of stalk with an at- 
tached slender peduncle bearing at the base the scar of 
a branch (Plate VI, D). Considering all of these speci- 
mens together we now reach the conclusion that the long 
husk probably enclosed a cluster of at least two ears, each 
of which had its own husks which were only slightly 
longer than the ears. The situation is illustrated in the 


[ 14 ] 


diagram in Fig. 8. The specimens on 
which it is based are illustrated in 
Plate VI. 

That this conclusion may be more 
nearly correct than the earlier one is 
supported by the fact that in our ge- 
netically reconstructed wild corn pro- 
duced by combining components of 
the tunicate locus with genes from 
various popcorn varieties, we have en- 
countered exactly this situation: clus- 
ters of small ears each enclosed in short 
husks and the entire cluster enclosed 
in long husks which flare open at ma- 
turity. 


Fic. 3. Diagrammatic longitudinal section based on 
specimens illustrated in Plate VI showing how the 
long husks may have enclosed a cluster of short ears, 
each enclosed in its own husks. Solid lines represent 
actual specimens; broken lines indicate what may 
have been the additional parts. 


"TASSELS 


Only one specimen of tassel, a fragment of a central 
spike, was found and this occurred in the uppermost 
level. Its spikelets have relatively long glumes. Since 
the specimens of tassels found in the 1948 excavation, 
although described, have not been illustrated, we are 


[15 ] 


/ 
' 
\ 
' 
! 
| 
\ 
I 
I 
! 
1 
( 
‘ 
! 
\ 
| 
1 
\ 


including a photograph of them in Plate VII. The 
spikelets on most of the fragments are smaller and more 
delicate than those of the majority of modern varieties 
but in other respects are similar. 


CONCLUSIONS 


1. The earliest maize from the second (1950) Bat Cave 
expedition is more primitive than any of the specimens 
turned up in the first (1948) expedition. 


2. Maize from the lower levels of the cave is definitely 
a popcorn. There are several popped kernels among the 
prehistoric remains and other prehistoric kernels proved 
to be still capable of popping after having their moisture 
content raised. 


3. The earliest maize is probably a form of pod corn. 
At least it has relatively long soft glumes partly enclos- 
ing the kernels which are borne on long rachillae. These 
are characteristics of pod corn. 


4. The maize from the lowest levels has brown peri- 
carp color and is related to the Mexican race, Chapalote. 
Brown pericarp color is presumably the primitive or 
‘‘wild’’ color. Brown pericarp is less frequent than red 
and colorless pericarp combined in the upper levels of 
the cave. 


5. The occurrence of variegated pericarp in the 86-48” 
level] shows that a mutation system similar to that involv- 
ing the ‘‘modulator’’ factor was in existence at an early 
stage in corn’s domestication. 


6. Clear-cut evidence of the introgression of teosinte 
or T'ripsacum or both appears in the 86-48” level of the 
cave and there is some evidence of introgression in the 
earlier 48-60” level. 


[ 16 ] 


7. There is an increase in diameter of the rachis of the 
cobs from the lower to the upper levels; this is accom- 
panied by an increase in kernel-row number and in the 
length and width of the kernels. There is a slight de- 
crease in kernel thickness. 


8. The remains of the husks and other parts of the 
husk systems suggest that the long husks found in one 
of the levels of the 1948 excavations enclosed not a single 
ear but a cluster of ears each enclosed in its own husk. 


9, Comparisons with the prehistoric maize of other 
sites suggest that the earliest maize from Bat Cave should 
be dated at not earlier than 2800 B.C. and perhaps 
several centuries later. 


LITERATURE CITED 


Antevs, KE. Geochronology of the deglacial and neothermal ages. 
Jour. Geol. 61: 195-230, 


Arnold, J. R. and W. FF. Libby. 1950. Radiocarbon dates. Univ. 
Chicago Inst. Nuclear Studies. 


Dick, H. W. 1965. Bat Cave, Catron County, New Mexico. School 
of American Research Monograph No. 27. Santa Fe. 


Emerson, R. A., G. W. Beadle, and A.C. Fraser. 1985. A summary 
of linkage studies in maize. Cornell Univ. Agr. Exp. Sta. Memoir 
180, 


Galinat, W.C. and J. H. Gunnerson. 19638. Spread of eight-rowed 
maize from the prehistoric Southwest. Bot. Mus. Leafl. Harvard 
Univ. 20: 117-160. 


+, P. C. Mangelsdorf, and L. Pierson, 1956. Estimates of teosinte 
introgression in archaeological maize. Bot. Mus. Leafl. Harvard 
Univ. 17: 101-124. 


-and Ruppe. 1961. Further archaeological evidence on the effects 
of teosinte introgression in the evolution of modern maize. Bot. 
Mus. Leafl. Harvard Univ. 19: 168-181. 


Goodman, M.M. 1965. The history and origin of maize. North 
Carolina Agri. Exp. Sta. Tech. Bull. No. 170. 


Libby, W. F. 1951. Radiocarbon dates I]. Science 114: 291-296. 


Mangelsdorf, P. C. 1958. The mutagenic etfect of hybridizing maize 
and teosinte. Cold Spring Harbor Symp. Quant. Biol. 23: 409— 
421. 

-and W.C. Galinat. 1964. The tunicate locus in maize dissected 


and reconstructed. Proc. Nat. Acad. Sei. 51: 147-150. 


and R. H. Lister. 1956. Archaeological evidence on the evolu- 
tion of maize in northwestern Mexico. Bot. Mus. Leafl. Harvard 
Univ. 17: 151-178. 


, R.S. MacNeish, and W. C. Galinat. 1956. Archaeological evi- 
dence on the diffusion and evolution of maize in northeastern Mex- 
ico. Bot. Mus. Leal. Harvard Univ. 17: 125-148. 


[ 18 | 


——, R.S. MacNeish, and W.C. Galinat. 1964. Domestication of 
corn. Science 143: 538-545. 


——, R. S. MacNeish, and W.C. Galinat. 1967. Prehistoric maize, 
teosinte, and Tripsacum from Tamaulipas, Mexico. Bot. Mus. 
Leafl. Harvard Univ. (in press). 


— and R. G. Reeves. 1939. The origin of Indian corn and its 
relatives. ‘'exas Agr. Exp. Sta. Bull. 574. 


—~——and R. G. Reeves. 1959. The origin of corn III. Modern races, 
the product of teosinte introgression. Bot. Mus. Leafl. Harvard 
Univ. 18: 389-411. 


~—— and C. ESmith. 1949. New archaeological evidence on evolu- 
tion of maize. Bot. Mus. Leafl. Harvard Univ. 13: 213-247. 


Sturtevant, E. L. 1894. Notes on maize. Bull. Torrey Bot. Club 21: 
319-343, 503-523. 


[19 ] 


Prare I. A and B. Two views of a segment of a 
disarticulating “‘tripsacoid’’ cob from the 24-36” 
level. The pedicellate member of each pair of spike- 
lets is staminate. The kernels have brown pericarp 
and are almost completely enclosed by the glumes. 
With respect to the pistillate spikelets, this is a 
four-rowed ear. C. Tripsacoid cob from the 36-48” 
level showing highly indurated tissues of the rachis. 
Approximately half of the pistillate spikelets are 
solitary. D. This specimen appears to be the pis- 
tillate basal part of a tassel branch or an un- 
branched tassel. E. A tripsacoid cob from the 36- 
48” level showing highly indurated tissues of the 
rachis, a whorled phyllotaxy, and a solitary spike- 
let. F. Fragment of a cob from the 36-48” level 
showing the almost square shaped cupules which 
are similar to those of the prehistoric wild corn 


from Tehuacan Valley. All photos * 3.7. 


[ 20 ] 


PLATE | 


Prare Il. A. A well preserved cob from the 48— 
60” level showing relatively long, soft lower and 
upper glumes and one cupule with a spikelet re- 
moved illustrating prominent cupule rims. This 
cob appears to be that of a form of pod corn. B. 
Fragment of an eroded cob from the 36-48” level 
showing a tendency for adjacent cupules to fuse 
and their rims to disappear. C and D. Two views 
of a fragment of an ear from the 36-48” level show- 
ing (1) the slender rachis and the long lower 
glumes and paleas and (2) the small round kernels 
with brown pericarp partially surrounded by the 


ri ~ 


glumes and other floral bracts. All photos ™“ 3.7. 


PLatreE II 


Piare Ill. A. Three cobs from the lowest level of 
the 1950 excavation. The diagrammatic longitudi- 
nal section illustrated in Fig. | is based on one of 
these. B. An evolutionary series of Chapalote-ty pe 
cobs, the shortest from the 48-60” level, the longest 
from the 12-24” level. C. Typical kernels with 
brown (left), red (center), and colorless pericarp. 
D. A prehistoric popped kernel from the 36-48” 
level compared with a popped kernel produced by 
exposing a prehistoric kernel from the same level 


to heat. All photos actual size. 


III 


PLATE 


Puate LY 


Typical cobs related to the Mexican race, Blando de Sonora from the 23-36” 


levels. 


A. Specimens of peduncles arranged in sequence from the lower to the upper levels. The vertical rows represent the 
36-48, 24-36/, 12-24’", 0-12” levels respectviely. B. Remains of husk system from the 36-48” level. Actual size. 


Prare VI. A. A long husk from level Il of the 
1948 excavation. B. A Chapalote-type cob from 
the 1948 collection. C. A husk from the 36-48” 
level which might have enclosed an ear of about 
the size represented by the cob in B. D. Piece of 
a stalk from the 36-48” level with peduncle of an 
ear arising fromit. At its base the peduncle has the 
sear of a branch. EK. A slender peduncle of about 
the diameter of the branch once attached to the 
base of D. These specimens are combined in a 
diagrammatic longitudinal section in Fig. 8. Three 


fourths actual size. 


VI 


Prat. 


PLATE VII 


| | | | | | | | | | 
cM. | 2 3 4 5 6 7 S§ 9 10 


Fragments of tassel branches from the 1948 excavations. The vertical rows from 


left to right represent levels VI, V, IV, 1V, IV respectively. 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY | 


CampripGre, Massacuuserrs, Decemper 27, 1967 VoL. 22, No. 2 


PREHISTORIC MAIZE, TEOSINTE, AND 
TRIPSACUM FROM TAMAULIPAS, MEXICO 


BY 
PauL C. ManGetsporr, RicHarp S. MacNEIsu, 
AND Wa.LTON C. GaLinat * 


IN an earlier paper (Mangelsdorf et al., 1956) we de- 
scribed the prehistoric maize uncovered in archaeological 
excavations conducted by MacNeish in 1949 in La Perra 
Cave in the state of Tamaulipas, northeastern Mexico. 
The earliest of this maize, dated at 2500 B.C. by radio- 
carbon determinations of associated wood and leaves, 
was identified as a precursor of a still existing Mexican 
race Nal-Tel, one of the four Mexican races described 
by Wellhausen et al. (1952) as Ancient Indigenous. We 
did not regard this maize, although primitive in some 
characteristics, as wild corn. 

While excavating La Perra Cave, which is located in 
eastern Tamaulipas, MacNeish also made some prelimi- 
nary soundings in several caves in southwestern T'amau- 
lipas which persuaded him that still earlier corn, perhaps 
even prehistoric wild corn, might be found in the lower 
levels of the refuse of these caves. Accordingly in 1954, 


* The authors are respectively : Fisher Professor of Natural History, 
Harvard University; Head Department of Archaeology, University 
of Calgary, Calgary, Alberta, Canada; Associate Professor, Waltham 
Field Station, University of Massachusetts, formerly Research Asso- 
ciate, Bussey Institution, Harvard University. 


[ 88] 


with the assistance of David Kelley, then a graduate 
student in anthropology at Harvard, he excavated two 
caves, Romero’sand Valenzuela’s, and sampled the refuse 
in a third, Ojo de Agua, all located in Infiernillo Can- 
yon. The earliest corn from these caves proved disap- 
pointingly to be not earlier than the La Perra Cave but 
slightly later, 2300-1800 B.C. It was, however, of a race 
different from the La Perra corn and showed some re- 
semblance to the early prehistoric corn of Bat Cave in 
New Mexico described by Mangelsdorf and Smith 
(1949). We now recognize both as being related to the 
prehistoric wild corn of Tehuacain Valley described by 
Mangelsdorf et al. (1964). 

Of even greater interest was the discovery in the Infier- 
nillo Canyon caves of several specimens each of teosinte, 
the closest relative of maize, and of Tripsacum, a more 
distant relative. The prehistoric remains of these three 
species of the American Maydeae are described below, 
following a brief description of the site and its environs. 


DEscRIPTION OF THE SITE 

The Canyon Infiernillo is located in southwest T'am- 
aulipas in the northern part of the municipio Ocampo. 
In terms of our previous investigations in Sierra de 
‘Tamaulipas (MacNeish, 1958; Mangelsdorf et al., 1956), 
this region is about 75 miles to the southwest and has a 
very different environment. Immediately southwest of 
the dry Sierra de Tamaulipas at an elevation never over 
1500 feet is the wide flat meandering Guayalejo River 
Valley with a tropical vegetation extending up from the 
south. West and southwest of this valley lie the first 
north-south oriented ridges of the Sierra Madre moun- 
tains. The lower slopes of these mountains are covered 
with a tropical, deciduous forest; from 2500 to 4000 feet 
the vegetation forms a cloud forest; and at higher eleva- 


[ 34 | 


tions a pine-oak forest occurs. Further to the west the 
valley becomes narrow and the ridges even higher. 
Covered with pine-oak forests at their summits, these 
ridges cast ‘‘rain shadows’”’ on the bottoms of the valleys 
which consequently are quite dry and have a xerophytic 
vegetation in which maguey, cactus, mesquite, and 
chaparral predominate. Canyon Infiernillo, to which we 
were guided in 1954 by Don Ignacio Guerra, is one of 
those dry canyons containing three caves named 
Romero’s, Valenzuela’s, and Ojo de Agua. These are 
all situated high above the canyon floor at the base of 
limestone cliffs and may have been the mouths of former 
underground rivers that have since become dry and in 
part filled in. These relatively deep tunnel-like caves are 
extremely dry and in prehistoric times would have been 
ideal places for living. Because of their dryness much of 
the refuse and remains of the ancient inhabitants has 
been almost perfectly preserved. 


The Stratigraphy 


All three caves had stratified occupational layers which 
not only revealed a long cultural sequence but also 
yielded many botanical specimens which could be 
brought to bear upon the problems of early agriculture 
and subsistence activities. 

Radiocarbon determinations and correlations of arti- 
facts with those of other sites permit us to recognize six 
more or less distinct cultural phases beginning at about 
2350 B.C. The characteristics of these phases have been 
described in detail elsewhere (MacNeish, 1958). Here 
we need be concerned only with their approximate ages. 
In sequence beginning with the most recent they are: 


San Antonio A.D. 1450-1800 
San Lorenzo A.D. 1050-1450 
Palmillas A.D. 200-800 


[ 35 | 


Mesa de Guaje 1200-400 B.C. 
Guerra 1850-1200 B.C. 
Flacco 2350-1850 B.C. 


The Specimens 


The maize specimens from only one of the three caves, 
Romero’s, have been analyzed in detail but an early cob 
from the Flacco phase of Valenzuela’s Cave has been 
included in the analysis as well as several specimens of 
teosinte and T'ripsacum. 

The collection of prehistoric remains of maize and its 
relatives described here comprises 12,014 specimens and 
includes virtually all parts of the plant: pieces of stalk, 
leaves, husks, cobs and cob fragments, tassel and tassel 
branches. There are a large number of quids of chewed 
stalks, young ears, and tassels as well as several speci- 
mens of teosinte and 7?7psacum. A brief description of 
the remains in the several categories follows. 


CLASSIFICATION OF THE Coss 


All of the intact cobs as well as those almost intact 
were classified on the basis of their resemblance to the 
existing races of corn in Mexico described by Wellhausen 
et al. (1952). Nine different races or subraces were iden- 
tified among the prehistoric cobs. Their relative fre- 
quency in terms of percentage of total cobs in the several 
levels of the cave is shown in Table I. 


The Chapalote Complex 


The great majority of the cobs, about two thirds of 
the total, were identified as belonging to the race Chap- 
alote or its precursors or derivatives. This race is found 
today only in western Mexico (Wellhausen et a/., 1952) 
but it was once much more widespread. The prehistoric 
wild corn uncovered in caves in the valley of Tehuacdn 


[ 36 ] 


Tape I, Classification of the Prehistoric Specimens of Maize and Its Relatives 
Associated with Six Cultura] Phases in Caves in Southwestern Tamaulipas. 


Cultural Phase 
8 3 © .: Ss) oO a 
: : ie) + S “= O ° oO 
Classification 7] <7 Si 8 bes faa eQ g 
of aS a vs ” 6|Fo6 S vs o 
Specimens Go) ce |e te el awe 7 = = 
qj .}/s .} gil Ea] ga ue 
eAleA|/#a}28]8S/ gs] 8 
Adi Hc(/ed/ sl] 5eiea| g 
Cobs 
Pre-Chapalote 2 14 1 17 
Early Chapalote 5 2 24 56 46 133 
Tripsacoid Chapalote 299 356 457 307 1ST 1546 
Chapalote 108 122 116 25 371 
Breve de Padilla 350 279 188 19 836 
Palomero Jalisciense T 15 11 4 37 
Harinoso de Ocho 19 10 3 32 
Nal-Tel 23 17 40 
Tuxpefio 3 3 
Unidentified Fragments 229 71 49 53 55 457 
Other Maize Parts 
Stalks 13 12 5 11 6 47 
Leaves 4 2 2 1 9 
Husks 45 60 70 42 2 219 
Tassels 6869 | 904 807 1s 4 8099 
Quids 44 39 63 5 151 
Maize Relatives 
Teosinte 1 7 1 9 
Maize-teosinte Hybrids 1 2 3 
Tripsacum 1 1 9 1 5 
Total Specimens 12,014 


[ 37 ] 


in southern Mexico is related to Chapalote (Mangelsdorf 
et al., 1964) as is also the earliest prehistoric corn from 
Swallow Cave in Sonora, Mexico (Mangelsdorf and 
Lister, 1956) and from Bat Cave in New Mexico 
(Mangelsdorf et a/., 1967). Ears, kernels, and cobs of 
this race have also been identified among the prehistoric 
specimens from archaeological sites in Arizona, Utah, 
Colorado, Nebraska, Oklahoma, and ‘Texas. This race, 
originating from its wild progenitor in southern Mexico, 
spread through western and northwestern Mexico and 
from there throughout the region now the southwestern 
United States. It has not been found, however, either 
archaeologically or among existing races, in lowland 
eastern Mexico. There the earliest corn is Nal-Tel, a 
race closely related to Chapalote and perhaps stemming 
from it. Nal-Tel is found today in Yucatan and Cam- 
peche and sporadically in Oaxaca and Guerrero. One 
collection of this race has been made in San Luis Potosi 
(Wellhausen et a/., 1952). The chief difference between 
these two ancient races is in pericarp color, Chapalote 
having brown and Nal-Tel orange pericarp. Other dif- 
ferences include shape of the ear and hairiness of the 
cupules. Although the differences are not consistent, 
Chapalote tends to have a longer, more tapering, cob 
than Nal-Tel and to have cupules which are less hairy. 
The fact that Chapalote is the predominating race in 
the prehistoric cobs from Canyon Infiernillo caves while 
the early corn from La Perra Cave is exclusively Nal-Tel 
(Mangelsdorf et al., 1956) suggests that races of culti- 
vated maize, like other artifacts, are characteristic of the 
cultures to which they belong. The people who occupied 
La Perra Cave were evidently related to lowland agri- 
culturists of eastern Mexico while those of the Canyon 
Infiernillo caves had their affinities westward and north- 
ward (MacNeish, 1958). It would also appear that races 


[ 38 ] 


of maize, although clearly related to each other such as 
Chapalote and Nal-Tel, had early become adapted to 
different highland and lowland environments and tended 
to spread within these ‘“‘ecological zones.’’ This in turn 
suggests that perhaps it was only at the latest prehistoric 
levels that the culture and agriculture of the highlands 
and lowlands merged to become the single ‘‘culture 
area’’ Meso-America. 

Pre-Chapalote. (Plate VIII, A). Following the no- 
menclature employed in our earlier descriptions of pre- 
historic maize (Mangelsdorf et al., 1956; Mangelsdorf 
and Lister, 1956) we have called the earliest corn un- 
covered in Romero’s Cave Pre-Chapalote. The cobs of 
this subrace have the same tapering shape as those of the 
modern Chapalote but are much smaller. None are as 
small, however, as the prehistoric wild corn from ‘Tehua- 
cin Valley. We assume that even the earliest corn from 
the Tamaulipas caves is cultivated corn. 

Pre-Chapalote makes its first appearance (a single cob) 
in the Flacco phase of Valenzuela’s Cave dated at 2850— 
1850 B.C. It appears last in the Mesa de Guaje phase 
dated at 1200-400 B.C. The total number of cobs of 
this subrace is 17. 

Early Chapalote. (Plate VIII, A). The cobs of this 
subrace, comprising 133 specimens, are intermediate in 
size between those of Pre-Chapalote and modern Chap- 
alote. In other respects the cobs are quite similar to 
those of modern Chapalote. Cobs of this subrace ap- 
peared first in the Guerra phase dated at 1850-1200 B.C. 
and last in the San Lorenzo phase, A.D. 1050-1450. 

Tripsacoid Chapalote. (Plate XI, A). Slightly more 
than half of all identified cobs, 1546 specimens, were 
assigned to a subrace which we called Tripsacoid Chap- 
alote. The cobs are quite similar to those of Chapalote 
but differ in the induration of their tissues, especially of 


[ 39 ] 


the rachises and glumes which are highly indurated. 
Cobs of this race are probably the product of hybridiza- 
tion of Chapalote with corn’s closest relative, teosinte. 
Specimens of teosinte and corn-teosinte hybrids were 
found among the prehistoric vegetal remains. 

The Tripsacoid Chapalote appeared first in the Guerra 
phase but at a later level than the Karly Chapalote. A 
single cob occurred in level 5 and the subrace was well 
established (126 cobs) in level 4b which represents the 
end of the Guerra phase probably about 1500-1200 B.C. 
This corn became the predominating type in the two 
succeeding phases, 1200 B.C.—A.D. 800, and thereafter 
was gradually replaced by other races, persisting, how- 
ever, aS a prominent component in the complex until 
A.D. 1800, after the arrival of the Spaniards. 

Chapalote. (Plate VIII, B). Cobs of a type quite 
similar to those of modern Chapalote appeared first in 
the Mesa de Guaje phase, 1200-400 B.C. They increased 
in frequency in the Palmillas phase (14.5 percent of all 
cobs) and continued to maintain approximately this fre- 
quency until the end of the series. A total of 361 cobs 
were assigned to this category. 

Wellhausen ef al. (1952) collected modern Chapalote 
in only two states in Mexico, Sinaloa and Sonora, but 
as pointed out above it must at one time have been 
much more widespread. 

Breve de Padilla, (Plate X, A). Making its first ap- 
pearance only slightly later than modern Chapalote— 
level 4a of the Mesa de Guaje phase—is a race called 
Breve de Padilla by Wellhausen (unpublished). 

The origin of this race is not definitely known. Its 
cobs are longer and thicker than those of Chapalote and 
it may be the product of hybridization between Chapa- 
lote and Harinoso de Ocho, a race originally from South 
America and still found sporadically in western Mexico. 


[ 40 ] 


Not altogether consistent with this suggestion is the fact 
that Breve de Padilla, the suspected hybrid, appears be- 
fore Harinoso de Ocho, the putative parent. However, 
the former occurs first in the upper level of the Mesa de 
Guaje phase, the latter in the lower level of Palmillas, 
the succeeding phase. Also, the Harinoso de Ocho had 
so low a frequency at all levels that its absence in the 
Mesa de Guaje phase may represent nothing more than 
a sampling error. 

Whatever its origin, Breve de Padilla appears to have 
been a productive race which rapidly replaced other 
races. Starting with an initial frequency of 4.6 percent, 
the cobs of this race increased in succeeding phases to 
23.5, 85.1, and 43.1 percent. In the San Antonio phase, 
A.D. 1450-1800, this was the predominating race. Alto- 
gether 836 cobs were assigned to this race. 

Cobs identified as those of Breve de Padilla had pre- 
viously been found among the later prehistoric specimens 
uncovered in the Sierra de Tamaulipas caves (Mangels- 
dorf et al., 1956; MacNeish, 1958) but their frequency 
was low. This may be regarded as a further bit of evi- 
dence that this race is western in origin and that La 
Perra and Canyon Infiernillo caves represent different 
peoples with respect to their agriculture. 


Minor Races 


A total of 112 cobs were found which could not be 
assigned to any of the races described above but all 
seemed to be related to others of the existing races de- 
scribed by Wellhausen et al. (1952). The following races 
were identified among the cobs. 

Harinoso de Ocho. (Plate XI, B). As already men- 
tioned Harinoso de Ocho first appeared in the lower level 
of the Palmillas phase, A.D. 200-850. At no time, how- 
ever, did it attain a high frequency and the total num- 


[ 41 ] 


ber of cobs assigned to this race was only 82, about one 
percent of the total identified cobs. 

Although not common in Mexico today, Harinoso is 
important as one of the progenitors of a number of races. 
According to Wellhausen et al. (1952) Harinoso de Ocho 
has given rise directly to the widely grown race Tablon- 
cillo and indirectly to Jala, Bolita, Celaya, and Cénico 
Norteno and is also related to Olotillo of southwestern 
Mexico. Evidence presented by Galinat and Gunnerson 
(1963) indicates that Harinoso de Ocho is the progenitor 
of the eight-rowed flour corn of the upper Mississippi 
and the eight-rowed flint corn of New England. This in 
turn makes it one of the ancestors of Corn Belt dent 
corn of the United States. It may also be, as suggested 
above, one of the progenitors of Breve de Padilla which 
became the predominating race of Romero’s Cave be- 
tween A.D. 1450-1800. 

Nal- Tel. (Plate X, B). A total of 40 cobs were as- 
signed to the race Nal-Tel which is the eastern low- 
land counterpart of Chapalote and was the predominating 
race in La Perra Cave in eastern ‘lamaulipas. Its low 
frequency in Infiernillo Canyon compared to the high 
frequency of Chapalote indicates that there was little 
interchange of cultivated corn varieties between the 
peoples represented by these two caves which are physi- 
cally only about 75 miles apart. 

Palomero Jalisciense. (Plate IX, B). A total of 37 
cobs were assigned to this race which is described by 
Wellhausen et a/.(1952) as a subrace of Palomero Tolu- 
queno, one of the Ancient Indigenous races of Mexico. 
The subrace differs from the ancestral race in having 
thicker cobs and a higher kernel-row number. Among 
the living races of Mexico, it has been collected only in 
southern Jalisco at elevations of 2600 to 2700 meters. 
In comparison to Palomero ‘Toluqueno, it is slightly 


[ 42 ] 


more vigorous, has a stronger root system, and is later 
in maturity. Also the ears are somewhat less tapering 
than those of Palomero Toluquefo. Wellhausen ef al. 
(1952) suggested that these modifications had probably 
been brought about through introgression of Oloton, a 
race common in the high altitudes of the state of Chiapas, 
Mexico, and in Guatemala. 

Tuxpeno. (Plate XI, A). Three unusually large cobs 
occurred which showed some resemblance to Tuxpeno, 
the predominating modern race of the lowlands of east- 
ern Mexico. There is some question, however, whether 
these cobs are actually of this race. A combination of 
higher than average hybrid vigor and better than average 
growing conditions could have produced unusually large 
cobs of the race Breve de Padilla, which could be con- 
fused with cobs of Tuxpeno. On the other hand, it is 
possible that 'Tuxpefo, which occurs in this part of 
Tamaulipas today, reached there in prehistoric times and 
it is to avoid overlooking this possibility that we include 
here a specific mention of these three large cobs. 


OTHER Parts OF THE CorRN PLANT 


A total of 8525 specimens of other parts of the corn 
plant were identified. These included stalks, leaves, 
husks, tassels, and chewed quids. 


Stalks 

Like the cobs, the pieces of stalks, 47 in number, 
showed an evolutionary sequence with respect to size, 
the earlier ones being on the average more slender than 
the later. 


Leaves 

Nine leaves or leaf fragments among the specimens 
add no significant information since they are similar to 
the leaves of modern corn. 


[ 43 ] 


Husks 
Like the cobs and stalks, the husks, a total of 219 


pieces, showed an evolutionary sequence with respect to 
size, the earlier ones being on the average shorter than 
the later ones. 

Tassels 

An amazingly large number of tassels, tassel branches, 
and tassel fragments, 8099 specimens in all, were found 
among the vegetal remains. There is great variation 
among these; the later ones are indistinguishable from 
those of modern races of corn; some of the earlier ones 
have smaller spikelets. Although some students of maize 
(Anderson and Cutler, 1942) consider the tassel to be an 
especially useful organ in classifying maize, we have not 
yet been able to discover any clear-cut, diagnostic char- 
acters which will allow us to assign the tassels, as we have 
the cobs, to recognized races. We are, however, preserv- 
ing all the specimens of tassels in the hope that some 
future student of prehistoric maize may see in them 
more than we have so far been able to discern. 

The most puzzling aspects of the tassels is why they 
should have been preserved at all. Perhaps young tassels 
still containing their anthers served as a source of food. 
The ancient peoples who occupied this cave seem to have 
led a precarious existence with respect to their food sup- 
ply and they chewed, presumably for the sugar they 
contained, a great variety of plants and plant parts. One 
of the specimens among the quids is undoubtedly that 
of the chewed young tassel since partly chewed staminate 
glumes can be identified in it. Also, we now know that 
corn pollen is rich not only in a number of amino acids 
but also in vitamins and minerals. Anthers filled with 
ripe pollen may be veritable little vitamin-mineral cap- 
sules and may have supplied some much needed elements 
to the diet. However, the majority of the prehistoric 


[ 44 ] 


tassel branches are those of matured tassels that have 
shed their pollen and lost their anthers. We can think of 
no use to which they might have been put for food or any 
other purpose except possibly a primitive ceremonial one. 


Chewed Quids 


The refuse contained a large number of chewed quids 
some of which, a total of 151, were identified as those of 
corn. Quids composed of chewed tassels have already 
been mentioned. ‘Two other types were recognized: 
those produced by chewing young ears enclosed in husks 
and those resulting from chewing stalks. Both were 
probably chewed more for their sweetness than for the 
few calories which they added to a none-too-adequate 
diet. After identifying two partly chewed young ears in 
the refuse of La Perra Cave some years ago, we chewed 
their modern counterparts and found them sweet. 

The majority of quids were those of chewed stalks. 
Chewing evidently began at one end of a piece of stalk 
and continued until the entire piece was thoroughly 
masticated and only the fiber remained. The process 
must sometimes have been interrupted for a few of the 
quids had unchewed sections of the stalk still attached 
producing a quid with a stem, a structure somewhat 
reminiscent of a modern lollypop (Plate XIII, D). 

The chewing of both stalks and young ears must have 
been at the expense of subsequent grain production. 
Perhaps in this stage of culture being provident had not 
yet become a virtue and was seldom practiced. 


TEOSINTE 


We identified nine specimens of teosinte and three of 
maize-teosinte hybrids. The earliest specimen, a frag- 
ment of fruit case, occurred in feces in one of the lower 
levels of the Guerra phase dated at 1850-1200 B.C. 


[ 45 ] 


Other specimens occurred in the two succeeding phases. 
Several of these comprised clusters of spikes (Plate XII, 
A). 

The specimens which we identified as maize-teosinte 
hybrids differed from those of teosinte in having thicker 
stalks and non-fragile rachises. The one illustrated in 
Plate XII, B is of particular interest in resembling spikes 
which we have produced by introducing one of the tuni- 
cate alleles into teosinte thereby replacing the highly in- 
durated lower glume of teosinte with the herbaceous 
glume of tunicate maize. When we first encountered 
this specimen we wondered briefly whether it might rep- 
resent the ancestral form of corn postulated by the late 
R. A. Emerson: a soft-shelled form of teosinte.* Be- 
cause these specimens were borne on thicker stalks than 
the clusters of teosinte we concluded that they were 
maize-teosinte hybrids. 

Wecannot be certain whether the specimens identified 
as maize-teosinte hybrids are F; hybrids or segregates 
appearing in subsequent generations. They resemble F 
hybrids in their non-shattering rachises but differ from 
typical modern F; hybrids in having single rather than 
paired spikelets. 

The presence in the caves of remains of teosinte is 
puzzling. The seeds of teosinte are nutritious, having a 
higher protein content than those of corn (Melhus ef al., 
1953), but are enclosed in hard bony shells from which 
they are difficult to remove. Although this can be done 
by popping if the moisture content of the seeds is right 
(Beadle, 1939) there is no evidence from the prehistoric 
remains that teosinte was used in this way. On the con- 
trary Dr. E. O. Callen, who has made a study of the 
feces from these caves, has found a number of teosinte 
fruits with their hard bony shells still undamaged or un- 


* In conversation. 


[ 46 ] 


changed. Since it is unlikely that the consumption of 
intact teosinte fruits provides any satisfaction or nutri- 
tional benefit to the consumer, there must have been 
some other reason for their use. Hernandez states that 
teosinte seeds are a cure for dysentery (Wilkes, 1967) 
and although it is doubtful that this is true, the inhabi- 
tants of the Infiernillo Canyon caves may well have be- 
heved it to be. Or did the Indians of this region perhaps 
practice the custom of planting teosinte in their corn 
fields to improve the corn as did those of western Mexico 
(Lumholz, 1902) or those of some parts of Guatemala 
(Melhus and Chamberlain, 1953)? The prehistoric speci- 
mens do not distinguish between the two possibilities. 
The finding of several fruits in a leather container in a 
pit extending down froma Palmillas floor suggests only 
that the fruits were regarded as having some value. The 
occurrence of maize-teosinte hybrids suggests that teo- 
sinte grew in or near the corn fields in prehistoric times 
although it is unknown in Tamaulipas today. 

The early occurrence in this site of teosinte and of 
tripsacoid maize, presumably the product of teosinte in- 
trogression, May raise questions regarding the hy pothesis 
of Mangelsdorf and Reeves (1939, 1959) that teosinte is 
a hybrid of maize and 7T’ripsacum which may have oc- 
curred after the cultivation of maize began. There is still 
earlier evidence of tripsacoid maize, although not of 
either teosinte or 7'ripsacum, inthe prehistoric cobs from 
caves in the valley of Tehuacan. In these sites, the trip- 
sacoid corn first appeared as a single cob in the Abejas 
phase, 3400-2300 B.C., and had become well established 
in the later Ajalpan phase, 1500-900 B.C. The earliest 
corn in both Tehuacan Valley and in ‘Tamaulipas is non- 
tripsacoid corn. ‘The tripsacoid corn appears soon after- 
ward and in Romero’s Cave at the same level as teosinte. 
These several findings, although consistent with the hy- 


[ 47 ] 


pothesis of the hybrid origin of teosinte, by no means 
prove its validity. 
‘TRIPSACUM 


Slightly less puzzling, since it still grows in 'Tamauli- 
pas, is the presence of 7'ripsacum in the refuse of the 
Canyon Infiernillo caves. Seeds of 7’7ripsacum, like those 
of teosinte, are nutritious but are difficult to remove from 
the bony shells in which they are enclosed. They are not 
especially promising as a source of food yet must some- 
times have been gathered for this purpose. Gilmore 
(1931) found fruits of 7’77:psacum in the prehistoric refuse 
of a cave in the Ozarks. 

Five specimens of 7'ripsacum were found in the refuse 
and these included both staminate and pistillate spikelets 
(Plate XIII, B). We were not able to make positive 
identification with respect to species although one of the 
spikes seemed to bear a somewhat closer resemblance to 
T’. zopilotense—which has not been collected in Tamau- 
lipas—than to 7. dactyloides which occurs there now. 


SUMMARY 


1. The vegetal remains uncovered in the refuse in 
three once inhabited caves in southwestern Tamaulipas, 
Mexico, are of particular interest in containing specimens 
of all three of the American Maydeae: maize, teosinte, 
and T'ripsacum as well as hybrids of maize and teosinte. 


2. The collection, comprising 12,014 specimens, in- 
cluded virtually all parts of the maize plant: stalks, 
leaves, husks, cobs, and tassels as well as chewed quids 
of various parts. 


3. The corn is predominantly of the race Chapalote; 
about two thirds of all of the identified cobs are assigned 
to this race and its several subraces. 


[ 48 ] 


4. The predominance of Chapalote and its subraces 
suggests that the prehistoric corn and the agriculture 
which they represent have stemmed directly from south- 
ern Mexico where the wild corn with which domestica- 
tion began was identified as a progenitor of the still 
existing Mexican race Chapalote. 


5. Beginning at about 1850-1450 B.C. there is evi- 
dence of the introgression of teosinte into corn. More 
than half of the identified cobs were classified as Tripsa- 
coid Chapalote. 

6. Several specimens of maize-teosinte hybrids furnish 


direct evidence of the hybridization of maize and teosinte. 


7. A new race, Breve de Padilla, which became the 
predominating race in the last phase, A.D. 1450-1800, 
may bea hybrid of Chapalote with the western Mexico 
race Harinoso de Ocho. 


[ 49 ] 


LITERATURE CITED 


Anderson, Kk. and H. C. Cutler. 1942. Races of Zea Mays: |. Their 
recognition and classification. Ann. Mo. Bot. Gard. 29: 69-88. 


Beadle, G. W. 1939. Teosinte and the origin of maize. Jour. Hered. 
380: 245-247. 


Galinat, W.C. and J. H. Gunnerson. 1963. Spread of eight-rowed 
maize from the prehistoric Southwest. Bot. Mus. Leafl. Harvard 
Univ. 20: 117-160. 


Gilmore, M. R. 1931. Vegetal remains of the Ozark Bluff-Dweller 
culture. Mich. Acad. Sci. Arts and Letters 14: 83-102, 


Lumholtz, C. 1902. Unknown Mexico. Charles Scribner’s Sons, New 


York. 


MacNeish, Richard S. 1958. Preliminary archaeological investigations 
in the Sierra de Tamaulipas, Mexico. Trans. Amer. Philos. Soc. 
48: Part 6. 


Mangelsdorf, P. C., H. W. Dick and J. Camara-Herniandez. 1967. 
Bat Cave revisited. Bot. Mus. Leafl. Harvard Univ. 22: 1-82. 


~and R. H. Lister. 1956. Archaeological evidence on the evolu- 
tion of maize in northwestern Mexico. Bot. Mus. Leafl. Harvard 
Univ. 17: 151-178. 


, R.S. MacNeish and W.C, Galinat. 1956. Archaeological evi- 
dence on the diffusion and evolution of maize in northeastern Mex- 
ico. Bot. Mus. Leatl, Harvard Univ. 17: 125-150. 


= -, RLS. MaeNeish and W. C. Galinat. 1964. Domestication of 
corn. Science 143: 538-545. 
— and R, G. Reeves. 1939. The origin of Indian corn and _ its 
relatives. Texas Agr. Exp. Sta. Bull, 574. 


— and R.G. Reeves. 1959. The origin of corn. III. Modern races, 
the product of teosinte introgression. Bot. Mus. Leafl. Harvard 
Univ. 18: 389-411. 


[ 50 | 


—— and C. E. Smith. 1949. New archaeological evidence on evolu- 
tion of maize. Bot. Mus. Leafl. Harvard Univ. 13: 213-247, 


Melhus, I. E., F. Aguirre and N.S. Scrimshaw. 1953. Observations 
on the nutritive value of teosinte. Science 117: 34-35. 


and I. M. Chamberlain. 1953. A preliminary study of teosinte 
in its region of origin. Iowa State Coll. Jour. Sci. 28: 139-164. 


Wellhausen, E. J., L. M. Roberts and E. Hernandez X. in collabo- 
ration with P. C. Mangelsdorf. 1952. Races of maize in Mexico. 
Bussey Institution, Harvard Univ. 


Wilkes, H. G. 1967. Teosinte: the closest relative of maize. Bussey 
Institution, Harvard Univ. 


[ 51 ] 


Prarke VIII. A. The two cobs (left) of Pre-Chapalote ; 
the lower from Guerra phase, 1850-1200 B.C., the 
upper from the Mesa de Guaje phase, 1200-400 B.C. 
The remaining four cobs are Early Chapalote. Both 
types are characterized by relatively soft glumes and 
rachis tissues. Actual size. B. Typical cobs of the race 
Chapalote from the Palmillas phase, A.D. 200-800. 
The stick inserted in one of the cobs may have served 
as a handle for holding the ear near fire to toast the 
kernels, Actual size. 


[ 52 


> VIII 


PLATE 


Prare IX. A. Cobs of Tripsacoid Chapalote from 
the upper level of the Guerra phase, ca. 1400-1200 
B.C. This maize, the predominating type in the 
cave, has indurated tissues of the rachises and lower 
glumes. When the other floral parts are lost the 
cobs with their stiff, often curved, lower glumes 
resemble a coarse file or rasp. The specimen at 
left still bears three kernels which have the brown 
pericarp color and round shape characteristic of 
Chapalote. Actual size. B. Cobs of Palomero Jalis- 
ciense, a popcorn race related to Palomero Tolu- 
queno, one of the four Ancient Indigenous races of 
Mexico. Palomero Jalisciense is known today only 


in the state of Jalisco. Actual size. 


PuatrE IX 


"OZIS [BNJOY “Saavo OM} BY} AQ payUasoidad Sain}z[Nd IY} UIIAJIq S2l}ZIIIVA UIOD JO BHULYOIIJU! 9]}}I] SPA B19} 
JY} SULBVOIPUL BAVD Ss OJIWIOY Ul a1vI SBM ‘SEdI[NEWIB] Uld4ysvd Ul BARD B1Iaq kB] UL 9UO Suljeuiwmopaid ayy ‘aoed sy], 
‘OOIXIJY Uloysvo fo spue[MO] ayy UL ayo[edeYyD jo yedsajUNOD ayy SI YOIyM [A] [WN aVI 9Y} 0} HSuLSuoOjaq se payljuapt 
SQOD “| *S9}8IS paul) 94} Ul JUIP yeIg UsOD pu yUIY puB[suY MAN puK ‘oolxay Ul BBs PUR OT[LOUO[Ge] Surpnyo 
-UI SooBl Ulopom jo Jaquinu ® jo Ai}s90UB 94} Ul payedloljied sey OdIxayY Us1d}saM UWI} 199}¥] OY], ‘OYOG ep osouePy 
pue sjoyedeyy) Sulzipuqsy jo yonpoid ay} aq ABUT YOIYA oR B EL[IPE IP 2AdIg Jo sqoo ayy aie AoUaNbaay ul puosag “VW 


a 


9 Gv € @ 


Z 


Prare XI. A. These cobs, of which there were 
only three, appeared in the San Lorenzo phase, 
A.D. 1050-1450. They are similar in their charac- 
teristics to those of the race Tuxpeno, one of the 
most productive of the modern races of Mexico, but 
they may be unusually large cobs of Breve de 
Padilla, the predominating race in the upper levels 
of the refuse. Actual size. B. These cobs may be of 
the race Harinoso de Ocho which is postulated to 
be the putative ancestor of Breve de Padilla, illus- 
trated in Plate X, and of several other productive 


races, 


if 
4 “4 


PLATE 


[19 | 


Tee 
ee 


a 


A. A cluster of teosinte spikes with a number of the fruits exposed to view. These, like the fruits 
of modern Mexican teosinte, are triangular in shape and in this respect differ from the fruits of 
Guatemalan teosinte which are trapzoidal. B. A spike identified as that of a maize-teosinte hy- 
brid. This differs from spikes of teosinte in having a solid rachis and herbaceous, instead of in- 
durated, lower glumes. Note that the stalk on which it is borne is thicker than the stalks of teosinte 
shown in Plate XII, A. Actual size. 


TIX #1v1q 


Prare XIII. A. Tassel branches from the San An- 
tonio phase, A.D. 1450-1800. All specimens are 
similar in their botanical characteristics to the 
staminate tassel branches of modern corn where 
the spikelets occur in pairs, one member sessile, 
the other pedicelled. Actual size. B. The pistillate 
part of a prehistoric spike of T’ripsacum and the 
staminate part of another spike. Why 7'ripsacum 
should have been collected and preserved in the 
save is puzzling. Actual size. C. Some of the cobs of 
Tripsacoid Chapalote are fragile like the spikes of 
teosinte and break up into disc-like segments (left) 
which resemble closely the segments of some de- 
rivatives of modern maize-teosinte hybrids (right). 
These prehistoric specimens furnish indirect evi- 
dence of the hybridizstion of maize and teosinte. 
Direct evidence of such hybridization is provided 
by the maize-teosinte hybrids illustrated in Plate 
XII, B. D. Quids, the product of chewing tassels 
(left, center) and a stalk (right). Young ears en- 
closed in husks were also sometimes chewed. 


Actual size. 


PLatE NIII 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


OVEMBER 22, 1968 


CATALOGUE OF INFRARED SPECTRA 
OF FOSSIL RESINS (AMBERS) 
I NORTH AND SOUTH AMERICA’ 
BY 
Jean H. LANGENHEIM AND Curt W. Beck’ 


Fossit resins are of interest to the mineralogist, palaeon- 
tologist, botanist, entomologist, chemist, and archaeolo- 
gist, but, because their study straddles these many fields, 
they have not received comprehensive attention from 
any quarter. During the 19th Century, mineralogists 
described and named well over 100 fossil resins, mostly 
of European origin, and botanists and entomologists 
studied plant and insect inclusions, primarily in Baltic 


' Grateful acknowledgment is made for funds to support this study 
from National Science Foundation grants GB-1312 and GB-2397 
at Harvard University and GS-739 and GP-4729 at Vassar College. 
Also financial assistance was provided to Professor Langenheim by 
the Radcliffe Institute for Independent Study and to both Professors 
Langenheim and Beck by the American Philosophical Society for the 
Advancement of Useful Knowledge. Mr. Gil Hillman, Mr. Anthony 
Placeres and Mr. Sumner Slavin assisted with preparation of the spec- 
tra and Mrs. Patty Shepard prepared the illustrations. Appreciation 
is likewise expressed to Professor E. S. Barghoorn for his continued 
interest and support of the project as well as his criticism of the manu- 
script and to Mr. James Doyle for his helpful discussions of Creta- 
ceous stratigraphy and floras. 

?Jean H. Langenheim, formerly a Research Associate in the Bo- 
tanical Museum, is now at the Division of Natural Sciences, University 
of California at Santa Cruz. Curt W. Beck is at the Department of 
Chemistry, Vassar College, Poughkeepsie, New York. 


[ 65] 


VoL. 22, No. 


amber. In the New World there are no fewer resins, 
but, since they have been studied by biologists rather 
than mineralogists, few have been given names. 

Even now, the only reasonably inclusive listing of fossil 
resin occurrences, restricted largely to the named varie- 
ties, are to be found in the back pages of standard works 
on mineralogy or mineral chemistry, such as those of 
Dana (1895), Hintze (1983) or Doelter and Leitmeier 
(1930). Their description by standard mineralogical 
procedures, which are ill suited to complex, largely 
non-crystalline, organic materials, does not provide an 
acceptable basis for classification. It is indeed no ex- 
aggeration to say that fossil resins have not yet been 
ordered into a classification system. Mineralogical com- 
pendia make broad distinctions between ‘‘families’’ con- 
taining or lacking succinic acid, sulfur or nitrogen, but 
these superficial criteria do not reflect any meaningful 
relationship in establishing a natural classification system. 

Paclt (1953) has suggested asystem ordered by geologic 
age and botanical origin. ‘he former criteria, unfortu- 
nately, have had to be based on meager and scattered 
published evidence. Most often, the botanical origin, 
which is based on associated fossil plant remains, is in- 
dicated without supporting evidence because of lack of 
available data in the literature. Botanical source unques- 
tionably is the most significant basis for classification of 
fossil resins, since they are plant products the chemical 
composition of which is genetically controlled (Mirov, 
1961; 1967). Also study of the botanical source of fossil 
resin through geologic time provides an opportunity to 
observe evidence of biochemical evolution within the 
populations that produce the resin. Resins from some 
populations appear to have a relatively stable compo- 
sition through millions of years, whereas others have 
changed considerably. The determination of the origin 


[ 66 | 


of resin, however, isa difficult task because of the changes 
which resins undergo in the process of fossilization. 
Resins polymerize with age, presumably by a free-radical 
mechanism. As a result, the fossil product is a macro- 
molecule of high molecular weight, largely insoluble, and 
not amenable to most of the usual methods of attack 
used in elucidating the structures of organic compounds. 

We have found infrared spectroscopy a useful technique 
in botanical investigations of both fossil and modern resins 
at Harvard University and in the classification of fossil 
resins for archaeological ends at Vassar College. We now 
hope to use our diverse interests and competences to 
undertake the large task of a systematic study of fossil 
resins. The goal is to classify them first relatively, 1.e. 
sorting them into groups of resins which share the same 
gross composition as revealed by their infrared spectra. 
Weeventually expect to determine their botanical source 
by comparison of the infrared spectra of recent and fossil 
resins corroborated wherever possible by paleobotanical 
data. Some results of this collaboration have been pub- 
lished previously (Langenheim and Beck, 1965). 

Infrared spectroscopy is useful particularly in com- 
paring fossil and recent resins because polymerization 
preserves all simple functional groups of the recent resin 
with the exception of carbon-carbon double bonds. Also 
skeletal frequencies are damped but not usually extin- 
guished completely. Thus, a similarity remains between 
the spectra of recent and fossil resins in that certain ab- 
sorption peaks can be matched one-to-one, although the 
intensity of these peaks is usually much weaker in the 
fossil resins than it is in the recent ones, especially at 
longer wave lengths. 

Over the past few years, we have prepared infrared 
spectra of well over a thousand fossil resins and several 
hundred present-day resins. It now seems appropriate 


[ 67 ] 


to make this large body of evidence accessible to inter- 
ested research workers in the form of a catalogue which 
will be divided into three sections: 1, North and South 
American; If, Europe; III, Asia and Africa. A fourth 
section probably will present spectra of resins from living 
plants. 

We have attempted to procure as many specimens as 
possible of each kind of fossil resin from the leading col- 
lections of the New and Old World. Samples used in 
this section were obtained from the Harvard University 
Botanical Museum Paleobotanical Collections; Harvard 
University Mineralogical Collections; Harvard Univer- 
sity Entomological Collections; Museum of Paleontolo- 
gy, University of California at Berkeley, United States 
National Museum (Smithsonian), New York Museum 
of Natural History and Museum fiir Naturkunde, Kast 
Berlin. 

Since infrared spectroscopy is not yet a method widely 
used by botanists and mineralogists, we will here include 
a brief discussion of the major absorption bands of the 
fossil resins and their significance. Infrared radiation is 
absorbed by matter to provide the necessary energy for 
the motion of atoms within a molecule. These motions 
may be relatively simple (stretching and bending of in- 
dividual bonds between two atoms) or they may be very 
complex (involving larger portions of the carbon skeleton 
of the molecule). The simpler motions predominantly 
lead to absorption at low wave lengths (2.5-8 uw; 4000 
to 1250 em:!). Because they are essentially independent 
of the rest of the molecule, their absorption bands are 
remarkably stable in position and are readily assigned to 
specific functional groups. Thus, the first absorption band 
of fossil resins at 2.9 » (8500 cm-!) is due to the stretch- 
ing of hydrogen-oxygen bonds; the bending motion of 
these bonds causes absorption at about 6.1 » (1650 em! ). 


[ 68 | 


Some of the hydrox] groups responsible for these two 
bands without doubt pre-exist in the resin, but others 
are the result of atmospheric water vapor taken up by 
the sample in the course of its preparation (Beck et al., 
1966). The group of more or less well resolved absorp- 
tions near 3.4 » (2950 cm-!) is due to the stretching of 
carbon-hydrogen bonds; the bending motions of these 
same bonds lead to absorption near 6.8 » (1470 cm-!) and 
7.25 pw (1880 cm-!), 

The remaining prominent band in the lower region is 
due to the stretching of carbon-oxygen double bonds. 
This so called carbonyl band usually occurs near 5.8 pu 
(1700 cm:!) in the fossil resins. All these absorption 
bands are almost uniformly typical of all fossil resins and, 
therefore, of limited diagnostic value. Only the position, 
intensity, and resolution of the carbonyl band show sig- 
nificant variation. 

The upper region of the spectra (8 to 16 w; 1250 to 
625 cm-!) is more difficult to interpret in terms of chemi- 
cal structure, but it is nevertheless more useful than the 
lower region because it shows greater variety. Of prime 
importance is the region between 8 and 10 p (1250 and 
1000 cm-!), where absorption is due to carbon-oxygen 
single bonds. Since these vibrations are substantially 
influenced by the carbon skeleton of the entire molecule, 
it is rarely possible to assign these bands to very specific 
structural features. However, as akind of fingerprint of 
a fossil resin, this upper region is the one that we most 
often rely upon to classify resins into groups which are 
not only recognizable as having similar basic structures, 
but which we can increasingly relate to recent resins, and 
thus obtain important evidence about the botanical origin 
of the fossil resins. 

Absorption bands above 10 » (1000 cm-!) are, in gener- 
al, still more difficult to assign and must therefore be used 


[ 69 ] 


mainly as fingerprints. But there are a few major excep- 
tions. Bending motions of hydrogen atoms attached to 
unsaturated carbon atoms lead to absorptions which are 
highly characteristic and which yield useful structural 
information. In the fossil resins, the most important of 
these is the sharp band near 11.8 » (885 em-!) which can 
be attributed to out-of-plane bending of the two hydro- 
gen atoms of a terminal methylene group. The presence 
of this band in the spectrum of a fossil resin is of particu- 
lar interest, because it is also a feature of a number of 
resin acids which have been isolated from recent resins 
(agathic acid, copalic acid, catavie acid, eperuic acid), 
The absence of this band in fossil resin spectra, however, 
must be interpreted with caution, since terminal methy!] 
groups are easily oxidized. The lack of an 11.3 p» (885 
em-!) band in the spectrum of a fossil resin may, there- 
fore, indicate either that the resin never had a terminal 
methylene group or that this group has been oxidized as a 
result of long exposure. The hydrogen atoms attached to 
aromatic rings also cause easily recognizable absorptions 
in the upper regions. ‘They are rare in fossil resins, but, 
when they occur, they permit immediate assignment of 
the resin to the few botanical sources characterized by 
aromatic components. 

All of the spectra were made from amber in the solid 
state which was dispersed in potassium bromide pellets 
(Langenheim and Beck, 1965; Beck et al., 1965). The 
Perkin Elmers Model 137 spectrophotometer was used 
for the samples reported in this section. 

The spectra curves reproduced here can, of course, be 
only a small fraction of those now recorded in both of 
our laboratories. ‘Io make the selection, we have com- 
pared all the spectra of each species or variety in order 
to choose a few spectra which best represent the group as 
a whole. In addition, we have included spectra of special 


(70 | 


interest since they show features not shared by all mem- 
bers of the group. 

We should like to warn researchers not to put too 
much weight on a single infrared spectrum of any fossil 
resin. Resins are mixtures of appreciable heterogeneity, 
and the small samples used to prepare spectra may well 
differ significantly in composition, even when taken from 
a single specimen. It is always advisable to run two or 
more spectra of a given specimen and to rely only on 
those features which are common to all or, at least, to 
most spectra. 

Relatively few of the North and South American am- 
bers have been characterized chemically or physically and 
assigned mineralogical species names, as numerous of the 
European ambers have been. In some instances, how- 
ever, paleobotanical remains of considerable significance 
in determining the botanical source of the amber have 
been reported in New World deposits. For this reason, 
botanical information regarding the kinds of trees that 
might have produced the amber, in addition to the man- 
ner in which the resin may have been produced, have 
received particular attention in this presentation. Con- 
fusion regarding stratigraphic data and, hence age as- 
signments of the amber-bearing strata, are discussed 
where relevant to an understanding of the rapid evolu- 
tionary changes which occurred in some of the possible 
resin-producing components of Cretaceous floras in par- 
ticular. Spectra of modern resins will be compared with 
those of ambers in some cases. Only general comments 
regarding the origin of the amber will be made, however, 
since this paper is designed primarily as a catalogue. 
Detailed discussion of botanical origin will be presented 
elsewhere. 


CRETACEOUS AMBER 


Amber from Cretaceous beds have been reported in 
considerable amounts from the Kuk River drainage of 
the Alaskan Arctic Coastal Plain, from the beach of 
Cedar Lake, Manitoba, Canada, and from numerous 
occurrences along the Atlantic Coastal Plain from Massa- 
chusetts to South Carolina. Also a small amount has 
been recorded from Baja, California, Mexico. Spectra 
have been made from specimens from the above locali- 
ties. Although amber has also been noted to occur in 
Cretaceous rocks from Hardin County, Tennessee (Ross, 
1956), Ellsworth County, Kansas (Schoewe, 1942), the 
Black Hills of South Dakota, Eagle Pass, Texas, Mt. 
Diablo, California, as well as other localities, samples of 
these have not been available as yet for analysis. 

The Alaskan amber will be discussed first because its 
stratigraphic relations and associated plant fossils have 
been investigated more intensively than for other Cre- 
taceous ambers. ‘These studies provide background for 
interpretation of the botanic source in terms of the in- 
frared spectra. 


Alaskan Amber 


Amber is widespread in Upper Cretaceous coal and 
carbonaceous shales along the Kuk River drainage (Kuk, 
Ketik and Kaolak Rivers) of the Alaskan Arctic Coastal 
Plain (Langenheim, Smiley, and Gray, 1960; Smiley, 
1966). Reworked amber occurs also ubiquitously in 
Pleistocene beds and along recent stream deposits in 
this region. Several lines of evidence indicate that these 
Cretaceous amber-bearing beds are of non-marine origin, 
and age determinations rest entirely on the abundant 
plant megafossils and microfossils. The older Kuk flora, 
characterized by ferns of varied kinds and a mixture of 
primitive and modern conifers, is considered Cenomanian 


[72 ] 


(earliest stage in the Upper Cretaceous) in age (Smiley, 
1966); only a few amber samples were found in these 
strata. More amber occurred in the Ketik beds to which 
Smiley assigns a Turonian age. These beds contain the 
earliest record of abundant and varied angiosperms and 
the dominant Ketik conifers are modern types referable 
to extant genera in such families as'Taxodiaceae (Sequoia, 
Sequoiadendron, ? Taxvodium and Juniperus), Pinaceae 
(Pinus, Picea and ? Larixv) and Taxaceae (Cephalotaxus 
and ¢ Torreya). Most of the amber occurred in associa- 
tion with the Kaolak flora which Smiley designates as 
Senonian in age. It is characterized by taxodiaceous 
conifers (Sequoia, Sequoiadendron and ? Taxodium) and 
¢ Torreya in the Taxaceae. Gray (Langenheim, ef al., 
1960) noted an abundance of conifer pollen in this flora 
with taxodiaceous pollen the dominant type. Pine and 
spruce pollen also occurred frequently, but the rarity of 
megafossils suggests that the latter conifers may have 
inhabited slopes or uplands some distance from lowland 
depositional sites. Langenheim et al. (1960) concluded 
that the source of this amber was taxodiaceous trees 
growing close to lakes, coastal swamps or other water 
bodies from the evidence: 1) the amber has always been 
found with abundant taxodiaceous remains, 2) the pina- 
ceous pollen was derived from upland sources. 

This Alaskan amber has been described neither chemi- 
cally nor physically and has not been given a mineral- 
ogical name. Some of the material is transparent, al- 
though much is opaque. The clear amber varies in color 
from light yellow to red, deep golden brown and almost 
black. The opaque material varied commonly from a 
beige to molasses color. Most of the transparent amber 
is crack-free and has a conchoidal to subconchoidal frac- 
ture, whereas the opaque amber is commonly granular. 
Langenheim et al. (1960) indicate that most clear pieces 


[ 73 ] 


do not contain inclusions, although small bubbles and/or 
small plant fragments are abundant in some specimens. 
Most unbroken specimens are small (one-eighth inch in 
diameter) teardrop masses, subcylindrical pieces up to 
one-fourth inch in diameter or irregularly mammillary 
blobs. The teardrops may have dropped from the trunk 
or branches, and the cylinders may have broken from 
hardened runnel of resin attached to the surface of a 
branch. All of these shapes tend to demonstrate resin 
hardened in the atmosphere. Several occurrences of am- 
ber in coal with woody texture, however, suggest that at 
least part of the amber was derived from resin within 
the trunk of tree and thus not in a position to trap in- 
sects or wind-blown plant debris. 

Twenty-two spectra were run of Alaskan amber rep- 
resenting samples varying in color and other physical 
properties. Nineteen specimens were from the Kaolak 
River and three from the Ketik River localities; none 
were available from the Kuk River. Most of these col- 
lections were from placer deposits, although at two locali- 
ties the amber was collected im situ. All of the material 
is from the collections of the Museum of Paleontology, 
University of California at Berkeley. Three patterns 
appeared among the 22 spectra run (Plates XIV and 
XV). The most typical pattern is Type II (H-382); 15 
spectra are of this kind. The spectra typical of amber from 
the Raritan formation from Kreischerville, Staten Is- 
land, New York, is similar to Alaska Type II in the 
carbon skeleton. Type I (H-318) is a more generalized 
pattern (i.e., more flattened with less pronounced bands) 
and is represented by three spectra. Type III (H-318 
on Plate XV) occurs only in two spectra but is quite 
similar in the carbon skeleton to that of the amber from 
the Upper Patapsco Formation from Washington, D.C. 
There are, however, differences between the pairs Alaska 


[ 74 ] 


Il and Kreischerville, on the one hand, and Alaska III 
and Washington, D.C., on the other, in the intensity and 
resolution of the carbonyl band. The 11.3 » (855 em-!) 
band is most pronounced in Alaska If and Washington, 
D.C. (H 173 on Plate XV). 

Spectra of resin from various living members of the 
‘Taxodiaceae and Pinaceae are being run presently to see 
if correlation can be established with these Alaskan types. 
The Taxodiaceae are of particular interest, since associ- 
ated plant remains with the Alaskan amber point to this 
possible origin. As yet, however, relation of this amber 
to either members of the Pinaceae or T'axodiaceae is not 
evident, but further investigation of the modern resins 
is necessary. 


Atlantic Coastal Plain Amber 


Amber for spectral analysis has been obtained from 
twelve localities along the Atlantic Coastal Plain from 
Massachusetts to South Carolina (Fig. 1). In most cases, 
the amber was reported to occur in Potomac, Raritan 
or Magothy beds. Controversy has existed regarding the 
age of these stratigraphic units (Spangler and Peterson, 
1950; Dorf, 1952; Steeves, 1959). Recent palynological 
studies, however, have essentially confirmed age assign- 
ments previously based on plant megafossils (Berry, 
1909, 1910, 191lla,b,c, 1914, 1916 et al.; Dorf, 1952). 

The Potomac Group is the basal unit of the Atlantic 
Coastal Plain Cretaceous sequence in Maryland, Dela- 
ware and Virginia. It typically underlies the Raritan 
and Magothy Formations in Maryland and Delaware 
but is absent to the northin New York, New Jersey and 
Massachusetts, where the Raritan is the oldest Cretaceous 
unit. The Potomac Group appears to be entirely Lower 
Cretaceous in age (Brenner, 1968), with the Patuxent 
Formation being Barremian, the Arundel Formation 


[ 75 ] 


I 
II 
III 


IV 


CRETACEOUS 


Alaska (Arctic Coastal Plain) . 
Manitoba, Canada (Cedar Lake) . . . 
Atlantic Coastal Plain 
1. Kreischerville, New York 
2, Martha’s Vineyard, Massachusetts 
3. Harrisonville, New Jersey 
4. Cliffwood, New Jersey . 
5. Bordentown, New Jersey 
6. Roebling, New Jersey . 
7. Kineora, New Jersey 
8. Pemberton, New Jersey . 
9. St. Georges, Delaware 
10. Washington, D.C. 
11. Magothy River, Maryland 
12, Charleston, South Carolina . 
Baja California, Mexico 


EARLY TERTIARY 


Seattle, Washington . 
Dominican Republic . 
Chiapas, Mexico 
Para, Brazil 


UNKNOWN AGE 


Guayaquil, Ecuador 
Girén, Colombia 
Medellin, Colombia 
Cafion Diablo, Arizona . 
Greenland 

Montana . 


Number of spectra for all localities are indicated. 


[ 76 ] 


. 30 


oO NO = 


\ 
Pf Partly pe’ 
© orXaduk = 5 


’ 
‘ 


& 


Ficure I. Location of samples of amber in North and South America 


from which infrared spectra were run (see explanation on opposite 
page). 


[77 ] 


being Aptian, and the Patapsco Formation being re- 
stricted to the Albian Stage. 

The Raritan Formation, on the other hand, is Upper 
Cretaceous (Cenomanian-Turonian Stages), according to 
the palynological investigations of Groot, Penny and 
Groot (1961) and Kimyai (1966). The overlying Mago- 
thy Formation is Turonian-Coniacian (Early Senonian) 
in age (Groot, Penny and Groot, 1961 and Stover, 1964). 

It is impossible to sort out the stratigraphic difficulties 
for most of the Atlantic Coastal Plain amber deposits 
from the available literature. It is most important to be 
aware of the age differences of these deposits, however, 
and possible confusion regarding them. The age is par- 
ticularly significant in regard to change in floras and 
seemingly rapid evolution of coniferous elements that 
could have been the amber producers. Although angio- 
sperms were rapidly evolving during this time, none ex- 
cept Liquidambar are known to have beenresin-producing 
plants. 


Washington, D.C. Amber 


A single sample of amber from the Upper Patapsco 
l’ormation was collected by J. Doyle from the corner of 
Branch Ave. and O Street SE, Washington, D.C. The 
amber came from a clay associated with a lignite bed at 
the top of typical Patapsco clays (Lower Cretaceous, 
Albian Stage) which are directly overlain by cross-bedded 
sands thought to be Upper Cretaceous by C.F. Withing- 
ton (Doyle, 1966). Doyle analyzed the plant microfossils 
from these beds and found a typical Upper Patapsco 
assemblage as established by Brenner (1953). Brenner’s 
palynological studies indicate that the gymnosperms of 
the Potomac forest included members of the Podocarpa- 
ceae, Araucariaceae (including Araucariacites australis 
Cook., which is comparable to modern Araucaria and 


[ 78 ] 


A gathis), Pinaceae and the Cupressaceae-Taxodiaceae 
complex. Particularly abundant in some localities is 
Inaperturopollenites dubius (Potonie and Venitz) Thom- 
son and Pflug, which is considered to be a member of 
either the Taxodiaceae or Cupressaceae. Brachyphyllum- 
Pagiophyllum represented other common coniferous 
families which are now extinct. Brenner further states 
that these Potomac forests were probably similar in 
character to the warm-temperate gymnosperm and fern 
forests of New Zealand today. In these forests, Podo- 
carpaceae and Araucariceae are the dominant trees with 
a luxuriant growth of ferns characterizing the understory. 
He adds, however, that the presence of cycads and 
schizeaceous plants suggests more tropical conditions 
than exist in present New Zealand forests. 

Two spectra were run from a single piece of amber 
from this O Street sample. The spectra are similar and 
represented by H173 (Plate XV). This spectrum re- 
sembles also Type III from Alaska (Plate XV). 


Kreischerville Amber 


Hollick (1905) reported considerable quantities of am- 
ber from a Cretaceous deposit at Kreischerville, Staten 
Island, New York. He stated that the beds were an 
eastward extension of the Amboy Clay and sand series 
of New Jersey and thus included in the Raritan forma- 
tion. He further continued that they were ‘‘middle 
Cretaceous in age and approximately equivalent of the 
Cenomanian of Europe. . .’’ Hollick and Jeffrey (1909) 
indicated that both the Raritan and Cliffwood (Magothy) 
Formations were present at Kreischerville but that only 
the Raritan beds here were plant-bearing. Unfortunate- 
ly, the stratigraphic position of the Kreischerville de- 
posits has not been investigated recently. 

The amber occurs in strata containing closely packed 


[ 79 ] 


masses of leaves, twigs and fragments of lignite and 
charred wood. Most of it was distributed irregularly in 
the matrix of relatively thick accumulations of commi- 
nuted lignite. The amber occurs commonly as small 
teardrops or in irregularly shaped fragments varying 
from the size of a pin head to a hickory nut. It is gener- 
ally transparent and yellow or reddish in color, although 
some is opaque and grayish white. 

In 1907, Gies’ chemical analysis indicated the follow- 
ing percentage elementary composition; C,78.5; H,9.5; 
S, 0.25; O, 11.75. On destructive distillation, succinic 
acid appeared to be formed and a considerable ‘‘quantity 
of volatile sulphide was evolved.’” Ash content was only 
0.1%. 

Because the amber at Kreischerville occurs in close 
association with leafy twigs of Sequoia heterophylla Vel. 
and S. Reichenbach (Gein.) Heer, Hollick thought that 
Sequoia might also be the source of this amber. He in- 
dicates, however, other coniferous remains found in the 
Kreischerville clay which might have contributed amber: 
Widdringtonites Reich (KKH.) Heer, Juniperus hynoides 
Heer, Dammara microlepsis Heer and Pinus sp. 

Jeffrey and Chrysler (1906) and Jeffrey (1907), on the 
other hand, studied the lignites from Kreischerville and 
concluded that Pityorylon (Pinaceous) was the only 
genus which contained amber. This ‘‘pitoxyloid lignite’’ 
contained masses of large pieces of amber; smaller frag- 
ments were contained in the amber-bearing strata. ‘‘The 
amber enclosed in the lignite appears both in translucent 
shining condition and in dull ochraceous modification. ”’ 
It not only occurs in ‘‘pockets or nuggets but also as fine 
yellow threads or streaks corresponding to the normal 
resin passages in the wood.”’ 

Thus, only both a possible taxodiaceous and pinaceous 
source have been suggested for the Kreischerville amber. 


[ 80 ] 


Six amber specimens from Kreischerville were run 
from Harvard Botanical Museum Paleobotanical Collec- 
tion #52812. All of the spectra were similar to that shown 
in H 821 (Plate XIV). Thus, the possibility of two 
sources did not become evident in the particular samples 
analyzed. The similarity of the spectra from Kreischer- 
ville and from the Koalak Riverin Alaska is noteworthy. 
Although stratigraphic data for the Kreischerville de- 
posit have not recently been critically assessed, it appears 
that the amber here is probably only slightly older (Ceno- 
manian) than the Koalak amber (Senonian). Although 
a taxodiaceous source is strongly indicated for at least 
part of the amber in both localities, infrared spectra of 
modern resins of representatives of this family have not 
as yet provided corroborative evidence. This has been 
true for the Pinaceae as well. 


Magothy River Amber 


Troost (1821) described amber associated with lignites 
at Cape Sable, Magothy River, Anne Arundel County, 
Maryland. These are the strata which later were desig- 
nated as the Magothy Formation (Little, 1917). Troost 
indicated that this amber occurs generally as grains ‘“from 
the size of a mustard seed to that of pieces from 4 to 5 
inches in diameter.’’ Most of it is opaque, varying from 
brown to yellowish grey; transparent specimens seldom 
are found. Also some of it is ‘‘earthy’’, occurring as 
friable, porous masses. 

Although Troost was interested in the kind of wood 
with which the amber was associated, he was unsuccess- 
ful in identifying the lignites. Knowlton (1896), how- 
ever, found at this locality a log which contained small 
pieces of amber. Poor preservation of wood and its ten- 
dency to disintegrate with exposure left its generic deter- 
mination open to question. Knowlton finally concluded, 


[ 81} 


however, that ‘‘as nearly as can be made out, the struc- 
ture is that of Sequora or Cupressinovylon as the wood 
is known in the fossil state.”” Henamed it Cupressinoxy- 
lon Bibbinsi, and suggested that this origin was supported 
by Fontaine’s recognition of 15 species of Sequoia from 
similar lignites found elsewhere in the Potomac Group. 

Hollick and Jeffrey (1909) concluded that most of the 
conifers found in Raritan strata, which had previously 
been referred to Podocarpaceae, Cupressaceae or Taxo- 
diaceae (Sequoiineae), belonged to the Araucariaceae. 
They recognized, however, the abietinious genera Prep- 
wus and Pinus. Penny (1947), on the other hand, sug- 
gested that some of the Raritan conifers that Hollick 
and Jeffrey considered aracauriaceous were taxodiaceous. 
For example, Sequoia Reichenbachi (Geinitz) Heer, which 
is synonymous with Geinitzia Reichenbachii (Geinitz) 
Holl. and Jeff., Penny considers to be essentially the 
same as Sequoia ambigua Heer. He suggests that these 
species, as well as others of Geinitzia, are taxodiaceous 
rather than araucariaceous. Araucarian remains, none- 
theless, are common in the Magothy flora of New Jersey 
and Delaware. 

Five spectra were run of specimens from USNM col- 
lection 772871. This amber was generally opaque with 
some color variation from beige to brown. Four of the 
spectra gave the pattern shown in H 861 (Plate XVI1). 
Another specimen from this same collection gave the 
typical Baltic Succinite pattern (Langenheim and Beck, 
1965). 

A taxodiaceous source is indicated from the wood in 
which the amber was enclosed, although the identifica- 
tion of this wood is tenuous. The spectral type of this 
amber (Plate XVIII) forms a family of amber including 
those from St. George, Delaware (H 357), Bordentown, 
N.J. (H 225), Cliffwood, N.J. (H 422) and Kincora, 


[ 82 | 


N.J. (H 367). This type is more closely related to the 
type from Manitoba, Canada (Plate XVI), than it is to 
that from Alaska and Kreischerville (Plates XIV, XV). 


Cliffwood Amber 


The lignitic sands and clays at Cliffwood Bluff, New 
Jersey, on Raritan Bay were first included in the Raritan 
Formation, although they also have been referred to the 
Matawan Formation (Hollick, 1897; Berry, 1905a). 
However, the lignites, and amber-bearing beds in par- 
ticular, were then considered to be in the Magothy For- 
mation. Berry (1905a) states that ‘‘a secondary feature 
of the Magothy Formation is the occurrence of amber, 
which is found in the form of globular and tear-shaped 
drops, disseminated in the lignite beds from Cliffwood 
bluff, in New Jersey, southwesterly to the type amber 
locality of the coastal plain, at Cape Sable, Md.’’ He 
indicates further that he has not found amber in pieces 
‘larger than a lima bean, although occasionally one 
hears from the foreman of the clay pits of much larger 
masses having been found.’’ Doyle (pers. comm., 1967) 
confirms the Magothy age of at least some of the Cliff 
wood lignites on the basis of his palynological studies. 

Berry (1905a,b) studied the plant megafossils from 
Cliffwood and described various coniferous remains. 
Among the araucarians were Dammara and Araucarites. 
Penny (1947) also reported Araucarioxylon with tracheal 
pitting similar to modern species of A gathis and Arau- 
caria here. Various species of Pinus, Picea and Pitoxylon 
are reported for the Pinaceae. Holden (1913) also listed 
Pityoxylon from this locality. In the Taxodiaceae, Berry 
reported various species of Sequoia, Geinitzia and Cun- 
ninghamites. 

One spectrum (Smithsonian Collection with no num- 
ber, collected by E.W. Berry from the Cliffwood Brick 


[ 83 ] 


Company) H 422 (Plate XN VIII) is similar to that of the 
Magothy River type. 


Martha's Vineyard Amber 


Amber from Gay Head, Martha’s Vineyard, Massa- 
chusetts, was collected by Clifford Kay, United States 
Geological Survey, Boston, Massachusetts. It had been 
reported previously from this locality by Finch (1824). 
Only a few small specimens have been found in a dark 
gray silt rich in fine plant debris, including seeds, cones, 
needles, leaf fragments and a variety of plant tissue ex- 
cellently preserved (Kay, pers. comm., 1966). Unfor- 
tunately ,these remains have not as yet been identified. 
Palynological evidence from these beds (J. Doyle, pers. 
comm., 1967) indicates their belonging to the Upper 
Raritan or Lower Magothy Formations. 

The spectrum (H 111, Plate X XI) is quite distinctive 
from any of the other ambers from Cretaceous strata. 
The intensity of absorption in the carbon-oxygen single 
bond region (8-10) may indicate a higher percentage of 
carbohydrates (as in a gum-resin). he presence of a dis- 
tinct absorption peak at 11.84 (885 cm-!) is also notable. 


Amber from Other Atlantic Coastal Plain Localities 


Amber was obtained from six other localities along 
the Atlantic Coastal Plain. Stratigraphic data for these 
localities are unknown to the authors. It is assumed, 
however, that the amber most likely occurs in Raritan 
or Magothy beds. 

Three spectra of ** Ambrosine’’ from Charleston, South 
Carolina (USNM collection R7317), were made. The 
spectral type represented by H 210 (Plate XVII) is 
typical of that of Baltic Succinite (Beck et al., 1964; 
1966; Langenheim and Beck, 1965). Shepard (1870) 
suggested that this amber originated from conifers from 


[ 84 ] 


the ‘‘pleiocene epoch.’’ Stratigraphic data are lacking, 
but Ambrosine has generally been considered from Cre- 
taceous Atlantic Coastal Plain deposits (probably from 
the Tuscaloosa Formation, which does not differ greatly 
in age from the Magothy, according to Groot, Penny 
and Groot (1961) ). Shepard described the original collec- 
tion as an irregular, oval-shaped mass the size of a man’s 
fist, yellow-brown externally but clove-brown inside. It 
melted to a clear yellow liquid at 460° F, giving con- 
siderable succinic acid before melting. On fusion, a dense 
yellow oil was volatilized, producing an odor similar to 
the resin of pines. It was largely soluble in turpentine, 
alcohol, ether and chloroform. The name ‘‘Ambrosine’’ 
was coined by compounding the two words ‘‘amber’’ 
and ‘‘rosin.”’ 

Three spectra were run from Kincora, Burlington 
County, New Jersey, from the United States National 
Museum Collection 5610, Spectrum H 867 (Plate X VIT) 
is similar to that typical of the Magothy River family of 
ambers. A variation of this spectrum with a band de- 
veloped at 12—12.3 (833-813 cm!) also occurs. 

A single spectrum was made of amber from St. 
Georges, Delaware (USNM #82552). Spectrum H 357 
(Plate XVIII) is generally similar to those from the 
Magothy River and Kincora, New Jersey, localities 
(Plate XVII). Three spectra were made of amber from 
Bordentown, N.J., (USGS collection #95367). All three 
were similar to that represented by H 225(Plate XVIII), 
although the 9.8 (1020 cm-!) band is sometimes more 
accentuated. This spectral type also belongs to the 
Magothy River family. 

Eleven spectra were made of amber from USNM col- 
lection R 7289 from Roebling, N.J. Considerable vari- 
ation in spectra occurs from specimens in this single 
collection. The general variation is shown in spectra H 


[ 85 | 


350, H 416, and H 420(Plate XIX). <All of this amber 
is opaque, with the color varying from light beige to 
streaks of almost black. The variation in the spectra ap- 
pear to be due largely to intensity and resolution of the 
bands. Roebling Type I resembles the Magothy River 
family of spectra. 

Amber from Harrisonville, N.J. (Gloucester Co.) was 
reported by Kunz (1883). It was found from 20 to 28 
feet under a green sand or marl! containing invertebrate 
fossils which were considered ‘‘to belong to the middle 
bed of the Upper Cretaceous series.’’ Kunz indicated 
that no analysis had been made of this amber, but he 
felt “‘the similarity in specific gravity, hardness and igni- 
tion leaves little doubt of its being true amber, or of its 
having been derived from a gum closely resembling that 
which is the source of the Baltic and other ambers.’”’ 

The spectrum H 409 (Plate XX) disproves this rela- 
tionship completely. The Harrisonville amber differs 
from almost all other fossil resins in that it lacks a well 
defined carbonyl group. Carbon-carbon unsaturation 
causes a sharp absorption band at 6.1 (1650 cm!) of 
such intensity that the broader hydroxy] absorption in 
this region fails to obscure it. The distinctive pair of 
bands at 13.84 and 14.34 (750 cm-! and 700 em:!) is 
clear evidence for the presence of mono-substituted ben- 
zenoid rings. ‘This indicates a composition similar to 
that of the European fossil resin siegburgite which has 
been related to Liquidambar by chemical studies in 
which cinnamic acid and styrene were isolated (Klinger 
and Pitschki, 1884). hese aromatic resins form a small 
but distinctive family of their own, which differ dramati- 
cally from the usual terpenoid resins which alone can be 
called ‘‘true amber.”’ 

The resin from Pemberton, N.J. (H 417, Plate XX) 
is of the same aromatic type as the Harrisonville sample 


[ 86 | 


with the exception of lacking the broad intense band 
which obscures the region between 8 and 1] (1250 and 
990 cm-!) in the Harrisonville sample. It also may well 
be due to inorganic impurities (silicates) which are 
characteristic of the Kuropean siegburgite. 


Canadian Amber 


Tyrrell (1890) investigated the occurrence of amber 
associated with fragments of decayed wood on the beach 
around the southwestern margin of Cedar Lake, near the 
mouth of the Saskatchewan River, Manitoba, Canada. 
He reported abundant amber along the beach for a mile, 
ina band thirty feet wide with a minimum depth of two 
feet. He further estimated that this deposit contained 
1,437,280 (!) pounds of amber. He also discovered amber 
occurring less extensively on shores of other lakes north 
and east of Cedar Lake. Because the amber has been 
deposited secondarily, the geologic age is uncertain. As 
Carpenter et al. (1988) indicated, however, Cedar Lake 
is fed by an easterly flowing river that drains through 
lignitic beds of Jate Cretaceous age in Alberta and south 
Saskatchewan. Thus, other amber-bearing deposits from 
which Cedar Lake amber could have originated are not 
evident. Carpenter et a/. point out that the insect in- 
clusions also provide support for a Cretaceous age. Evi- 
dence for a more specific time during the long Cretaceous 
interval, however, is not available. 

Although it cannot be definitely related to the Cedar 
Lake deposits, amber with insects found recently at 
Medicine Hat, Alberta (Richards, 1966), is of some in- 
terest in regard to the Manitoban material. ‘The Medi- 
cine Hat amber is associated with lignitic deposits from 
the Foremost Formation (Belly River Series, Upper 
Cretaceous). The Belly River Series falls within the 
Campanian Stage, as K/Ar dates for the Bearpaw shales 


[ 87 ] 


that occur immediately above the Foremost horizons at 
Medicine Hat range from 72 to 73 million years. None 
of this amber has as yet been available for analysis. 
Amber collected by ‘Tyrrell was studied by Harring- 
ton (1891) and named chemawinite. Klebs (1896), 
apparently unaware that Harrington had described it, 
analyzed the same type of amber from Cedar Lake and 
called it cedarite. Schmid (1980), in his review of fossil 
resins, classifies them as (a) ‘‘ Bernstein which yields suc- 
cinic acid on distillation, the principal type being succin- 
ite, (b) those resembling Bernstein, including cedarite, 
and (c) those quite different from (fern stehende) Bern- 
stein, including chemawinite.’” Walker has already 
pointed out, ‘‘the two Canadian minerals, cedarite and 
chemawinite, are identical, as appears probable both from 
the chemical and physical data recorded by Harrington 
and Klebs, and from the localities from which the ma- 
terial was obtained.’” Harrington (1891) made the most 
complete chemical analysis of this amber. ‘The hardness 
was 2.5; the specific gravity 1.055 at 20° C. The mean 
percentages of constituents from elemental analyses 
were: C, 79.96% ; H, 10.46% ; O, 9.49% and ash 0.9%. 
The ash contained silica, alumina, iron, calecilum and 
magnesium. 20.01% of the amber dissolved in absolute 
alcohol and 24.84% in absolute ether. Small fragments 
heated in a closed tube began to soften at about 150° C. 
Heated to 800° C, the resin did not melt into a flowing 
liquid but had become soft and elastic and darkened 
from partial decomposition. Harrington concluded that 
this Canadian amber was more resistant to heat than 
Baltic amber. Succinic acid, so characteristic of Baltic 
amber (Succinite), is absent. These facts, in addition to 
the presence of more carbon and less oxygen than Baltic 
Succinite, led him to conclude that the Canadian amber 
‘is closer to Walchowite and to some of the more recent 


copals from India. ”’ 
[ 88 ] 


This Canadian amber is transparent, varying from pale 
yellow, red to dark amber in color. Walker (1984) re- 
ported that the smallness of the amber grains (58% being 
less than eight mesh and only 3% larger than two mesh) 
was a point of difference from that of the generally larger 
masses characteristic of Baltic amber. He suggested that 
this might indicate conifers of types yielding little resin 
when wounded, in contrast to those which produced 
resin copiously in the Baltic forests. 

Eleven spectra were run on amber from Cedar Lake. 
Ten of these spectra were of specimens from the Harvard 
University Entomological Collections. All of these gave 
the same pattern represented by H 316 (Plate XVI), 
although some spectra were sharper than others. Also 
the pattern of amber from Cedar Lake, Manitoba, is 
similar to Ambrite (H 226, Plate XVI) from New Zea- 
land (USNM #7312). Both of these spectra, in turn, are 
similar to resin from A gathis australis, the Kauri pine 
(H 265, Plate XVI). Since 4. australis is noted for its 
capacity to produce large quantities of resin, it is a logi- 
cal source for fossilized resin from New Zealand. A 
possible Araucarian source, such as 4 gathis or related 
araucarians, for this Canadian amber is interesting and 
suggestive. More resins of members of the Araucariaceae 
need to be analyzed, before a definite correlation is made. 
Also the possible presence of A gathis australis (called 
Dammura australis Lamb.) in the Kreischerville, N.Y., 
deposits is noteworthy. Certainly, in the Potomac (AI- 
bian), Raritan (Cenomanian-Turonian) and Magothy 
(Early Cenomanian) Formations along the Atlantic 
Coastal Plain, members of the Araucariaceae were com- 
mon, and an araucarian-type resin could well be ex- 
pected. Araucarian leaf remains, i.e., 47aucarites longi- 
folia, are reported from Upper Cretaceous beds in the 
Rocky Mountains (Lance) by Dorf (1942), although 


[ 89 ] 


remains similar to Agathis have not been listed. 

One Cedar Lake specimen (USNM #97080) gave a 
spectrum similar to that of Baltic Succinite (Langenheim 
and Beck, 1965). The presence of this spectral type 
might suggest two different botanical sources of the am- 
ber from the Manitoban deposit. 


Baja California Amber 


Buddhue (1935) described ‘‘Bacalite’’ as amber pre- 
sumably from Baja California, Mexico. Since this amber 
cannot be shown to have come from Baja California, 
Langenheim, Buddhue and Jalinek (1965) recognized 
that ‘‘it may have come from any place in the world.”’ 
They mention, however, the particular likelihood of its 
being from the amber deposits of Chiapas, Mexico. 

Hurd, Smith and Durham (1962) reported docu- 
mented occurrences of amber from Baja California, based 
on the discovery by A.C. Allison and F.H. Kelmer. 
Langenheim, Buddhue and Jelinek (1965) further elabo- 
rated on the age and occurrence of this amber. It has 
been observed at three localities between Punta San José 
and Punta Cabras. At each locality, a single nodule of 
amber from 14—2 inches in diameter was collected from 
siltstone or sandstone thought to be latest Campanian or 
sarliest Maestrichtian in age. At Punta Baja, about 10 
miles southwest of KE] Rosario, small fragments of amber 
occur in sandstone with carbonized leaf impressions. 
Metaplefenticiras pacificum Smith occurs in the same bed 
and indicates late Campanian age. Small grains of amber 
and carbonized wood were also reported from a sandstone 
interbedded with a claystone south of the mouth of 
Arroyo Rosario. 

Four spectra were run from a single collection (Uni- 
versity of California, Museum of Paleontology, B-8006) 
from the Punta San José locality. The material generally 


[ 90 | 


is Opaque, varying in color from beige to molasses to 
almost black. More variation occurred in the spectra from 
these few pieces than is frequently the case where a larger 
number of samples have been run, i.e., from Chiapas, 
Mexico and Alaska. Three spectra showing this variation 
are presented in Plate XXII. Type I (H 45) and Type 
II (H 820) are similar, but Type III differs in having 
a broader and more intense absorption band between 
9-10 (1110-1000 cm-!). This may be due to inclusions 
of siliceous material. Type III was taken from a light 
beige portion of a banded specimen of which Type II 
represents the darker portions. As indicated with the 
Roebling, N.J. amber, more heterogeneity must be ex- 
pected in the opaque amber types than in the trans- 
parent ones. 


Earty TerriAry AMBER 


Amber is reported to occur in strata from early ‘Terti- 
ary time(Kocene to Miocene) in various areas throughout 
North and South America. A fairly extensive deposit 
has been discovered in Chiapas, Mexico, and is being 
intensively investigated by geologists, entomologists and 
botanists. Thus, the Chiapas deposit can serve as a useful 
standard for comparison of other Tertiary ambers as the 
Alaskan deposit does for Cretaceous ambers. Small 
amounts of Eo-Oligocene amber occur near Seattle, 
Washington; amber of probable Oligocene age occurs 
in the Dominican Republic and of Miocene age in Para, 
Brazil. The age of amber from several other localities 
probably is early Tertiary, but stratigraphic data are 
lacking. 

Chiapas Amber 

Amber from Chiapas, the southernmost state in Mex- 
ico, has been reported in mineralogical studies (Helm, 
1891; Kunz, 1903; Tschirch, 1906; Hintze, 19338; 


[ 91 | 


Buddhue, 1935; Tschirch and Stock, 1936), but it has 
neither been analyzed chemically and physically nor been 
given a mineralogical name. The geologic occurrence 
was first recorded by Bose in 1905. A coordinated scien- 
tific investigation was initiated by entomologists and 
later included geologists and botanists (Hurd and Smith, 
1957; Hurd, Smith and Durham, 1962; Langenheim, 
1964, 1966, 1967; Langenheim, Hackner, and Bartlett, 
1967). 

The amber-bearing beds are ina sequence of primarily 
marine, caleareous sandstones and siltstones which are 
mostly exposed in landslides. All amber-bearing locali- 
ties are considered latest Oligocene to earliest Miocene 
in age, based on marine invertebrate fossils (Licari, 1960; 
Hurd, Smith and Durham, 1962). Pollen from the strata 
either containing the amber or immediately adjacent to 
them indicates development of mangrove vegetation of 
considerable complexity at or close to the site of depo- 
sition of the amber (Langenheim, Hackner and Bartlett, 
1967). 

Previous discussions have presented evidence support- 
ing the conclusion that this amber was derived from the 
leguminous genus Hymenaea (Langenheim, 1963; Lan- 
genheim and Beck, 1965; Langenheim, 1966; and 
Langenheim, 1967). Spectra were made of various resins 
from populations existing in Chiapas today. Each genus, 
as well as certain species populations, was clearly dis- 
tinghishable. Remarkably consistent spectra were ob- 
tained from 35 samples of Hymenaea Courbaril L. from 
various populations in Mexico, Guatemala, Dominica, 
Costa Rica, Guyana, Venezuela, Brazil and Ecuador. 
Forty specimens of Chiapas amber from the University 
of California, Museum of Paleontology Collections from 
various localities were chosen for spectra because they 
showed some difference in physical character. The color 


[92] 


of this amber varies from yellow, golden brown, red to 
black; the most common being the yellow and golden 
brown hues. Variety in appearance tended to indicate 
that several kinds of trees might have produced the am- 
ber. All but two specimens fit into two spectral patterns 
indicated as Type I (H_ 1A) and Type II (H 88), Plate 
XXIII. A third type (H 1C) gives an ill defined spec- 
trum. It was, therefore, initially assumed that these three 
patterns might indicate three different sources (Langen- 
heim, 1963). The two major types (14 samples of each), 
however, are similar to each other, except for the more 
general flattening of the bands and loss of the sharp band 
at 11.2—11.3p (895-885 cm-!) in Type II. The similarity 
is now thought to be sufficient to suggest that both Type 
I and Type II were derived from the same source and 
that the differences may be explained by additional at- 
mospheric oxidation and progressive polymerization. The 
two specimens representing Type III (H 1C, Plate 
X XIII) are black in general appearance, although small 
pieces show aruby red color in transmitted light. The 
pieces are extremely soft and friable, indicating that they 
had been partially decomposed. Type I had aspectrum 
similar to that of Hymenaea Courbaril (H 1B-IV, H 58, 
Plate XXVIII), although there are some differences. 
Langenheim and Beck (1965) explained that the broad 
band at 14.2-14.4 (705-700 cm!) due to unassigned 
skeletal vibration has disappeared in the amber. The 
11.3 (885 cm-!) band which can be assigned to =CHg 
out-of-plane deformation of terminal carbon-carbon 
bonds decreases, as would be an expected result of pro- 
gressive oxidation by atmospheric oxygen. The attend- 
ant formation of new carbon-oxygen bonds would also 
account for the slight shift of absorption maximum near 
8 (1250 em-!) which is more often at 7.6 (1295 em:!) 
in the Hymenaea resin but more often at 8. 1p (1285 cm!) 


[ 93 ] 


in the amber. It is also significant that none of the other 
resin producers in Chiapas today had spectra resembling 
the amber (Langenheim and Beck, 1965). Thus, the in- 
frared spectra have given convincing evidence that the 
Chiapas amber was derived from an ancestral member 
of Hymenaea. 

The presence of Hymenaea leaflets in the amber pro- 
vides further corroboration for the source (Langenheim, 
1966). This conclusion receives additional support from 
studies of the manner of resin production and accumula- 
tion in Hymenaea Courbaril (Langenheim, 1967) and the 
depositional conditions of the amber. HY. Courbaril pro- 
duces large quantities of resin that accumulate in the soil 
around the roots, and in Mexico today it grows common- 
ly along rivers that enter the ocean in mangrove-fringed 
estuaries. Analysis of the pollen in the amber-bearing 
beds indicates that the amber was deposited in such an 
estuarine environment with abundant mangrove present 
(Langenheim, Hackner and Bartlett, 1967). 


Seattle Amber 


Amber from the Renton Formation near Seattle, 
Washington, was discovered by J. Wolfe. Vine (1962) 
considers the Renton to be late Eocene to early Oligo- 
cene in age. Wolfe (pers. com., 1966) reports that there 
are no plant fossils at the amber locality itself, although 
they are common in the Puget Group, of which the Ren- 
ton Formation is a member. Conifers present are taxo- 
diaceous, including Sequoia, Metasequoia and Glypto- 
strobus. The dicotyledenous flora is rich but still largely 
unidentified. The La Porte and Comstock floras are 
about the same age as the Renton. 

Five spectra were run of amber from the Museum of 
Paleontology, University of California Collection B8427. 
Most of this amber occurs in small friable pieces. It is 


[ 94 ] 


pale yellow and transparent. Chemical and physical 
analyses have not been made. There was some variation 
in the spectral patterns (Plate X NIV) as indicated in 
Type land Type II (Spectra H 823 and H 42A). These 
spectral types from Seattle are similar to those of Types 
I and II from Baja California (Plates XXII and XXIV) 
which indicates possibly that they may be from the same 
source. There are no present suggestions for the botanic 
source from these two localities either from associated 
plant remains or from spectra of modern resins. 


Dominican Republic Amber 


Sanderson and Farr (1960) state that amber was first 
reported from what now is the Dominican Republic by 
Christopher Columbus during his second voyage to the 
West Indies between 1494 and 1496. No further refer- 
ence to Dominican amber during the next 400 years 
appears to have been recorded. In 1905, Sample described 
an amber-bearing formation in the Monte Cristi Range 
(Cordillera Septentrional). Lengweiler (1989) then re- 
ported that fragments of lignite, leaves, and insects, 
such as mosquitos and ants, were found in the amber. 

Sanderson and Farr (1960) indicate that the amber- 
bearing formation in the Dominican Republic are located 
at two principal sites in the Cordillera Septentrional 
north of Santiago between Altamira and Conca. The 
original site is the Pena (Tamboril) region in the two 
gorges of the Arroyo Capancho tributary of the Rio 
Gurado. The second site is below Pico Diego de Ocampo 
in the Palo Alto de la Cumbre region. Sanderson and 
Farr collected the amber in a light brown-dark grey, 
fine-grained, micaceous sandstone occurring below a silty 
shale at this second site. Brower (Sanderson and Farr, 
1960) believes this amber to be Oligocene in age, al- 
though the exact date remains in doubt. The amber 


[ 95 ] 


occurs irregularly as small broken fragments to large 
unbroken pieces. Transparent, unfractured pieces are 
common, but many small broken pieces are brittle and 
fracture. ‘The color varies from clear crystal through 
various amber-shades to deep red. Chemical or physical 
analyses have not been made of this amber. 

Six spectra were made of amber from the Dominican 
Republic. Type I (H 37) is from a specimen from the 
Palo Alto de la Cumbre site collected by Sanderson 
(Plate X XV). Three others from an unidentified locality 
near Santiago gave a pattern similar to this one from the 
known site. ‘wo were run of specimens from USNM 
collection #R6783, from Palo Quemado, Santiago Pro- 
vince, Santo Domingo. ‘These samples gave a more 
generalized spectrum as represented by (H 427) than the 
other four (Plate X XV). They are designated as spec- 
tral ype II, and represent probably more oxidized 
specimens as indicated by the almost complete loss of 
the 11.34 (885 cm!) band. The spectral pattern is unique 
for North and South American ambers examined thus 
far. No extant groups have given a pattern similar to 
this, and, with correlative paleobotanical data lacking, 
there presently is no hint as to botanical source. 


Brazilian Amber 


A single piece of amber has been collected from the 
Pirabas Formation near Capanéma, Para, Brazil. This 
formation is considered early Miocene in age and is ac- 
companied by numerous dicotyledenous leaves (Lelia 
Duarte, pers. com., 1966). Spectrum H 452 is closely 
similar to resin from extant Hymenaea Courbaril (Plate 
XXVIII), especially from Brazil (H 61) and Costa Rica 
(H 252). Hymenaea thus appears to have been the source 
of this amber as well as that from Chiapas, Mexico. 
Perhaps future paleobotanical studies on these beds will 
give additional supporting evidence. 


[ 96 ] 


AMBER OF UNKNOWN AGE 


The geologic age and paleobotanic information are not 
available for a number of amber collections. ‘These col- 
lections are being presented separately, although they 
will be related to ambers of known age wherever this 
appears justifiable. 


Greenland Amber 


‘T'wo spectra were made of amber from Hare Island in 
Greenland from a collection from Museum fiir Natur- 
kunde, Berlin. The amber occurred as droplets in coal. 
The spectrum (H 165) is poorly developed, due probably 
to oxidation (Plate XV). This amber is assumed to be 
Cretaceous, because of the extensive, well known Cre- 
taceous plant fossil localities in this area of Greenland, 
although there is no direct confirmation of this age. 


Montana Amber 


Nine spectra were run of the amber from Montana. 
Collections were available from the United States 
National Museum and the New York Museum of Natu- 
ral History. No geologic age is indicated for this amber. 
Four spectra were made from the USNM Collection 
#112822, listed only from ‘‘Montana.’’ They are repre- 
sented by the Type I pattern represented by H 404(Plate 
XX). This is the third example of the aromatic series de- 
scribed above from Pemberton, N.J. (H 417) and Har- 
risonville, N.J.(H 409). The Montana sample is virtually 
indistinguishable from that from Pemberton. Both lack 
the broad absorption band of the Harrisonville sample 
between 9 and 10p (1110-1000 em-!) which we have as- 
signed to silicate impurities. 

Five spectra were run from the N.Y. Museum of 
Natural History collection £18395, listed from Washoe, 
Montana. Four spectra gave the pattern of Type II 


[ 97 ] 


(H 405) and one gives Type III (H 425) shown in Plate 
X XI. The small differences between these two types 
may be due to oxidation and progressive polymerization. 
Both are similar to the Magothy River family. 


Cation Diablo, Arizona Amber 


One spectrum (H 865) of amber was run from USNM 
collection #62998 (Plate XXIV). The age of this sample 
is unknown; in fact, no data are available for it. The 
spectrum is somewhat like those belonging to the Atlan- 
tic Coastal Plain Magothy family of ambers. 


Keuadorian Amber 


Amber from a “‘large deposit’” near Guayaquil, Kcua- 
dor, was reported by Johnston (1888) and named Guaya- 
quillite. No geologic age was given. ‘Two kinds, a light 
yellow, almost homogeneous material and a dark brown, 
bituminous-like substance, were found. The ‘‘pure”’ 
mineral is opaque, powders easily, and is soluble in alco- 
hol. The specific gravity is 1.092 and the melting point 
between 157°-212° C. 

Three spectra were made from USNM collection 
/R7328. Spectrum (H 212) is typical for the three (Plate 
X XV). Frondell (1967) has shown that X-ray diffraction 
patterns of Guayaquillite are similar to those of a sample 
of various species of Protiwm, a genus in the Burseraceae. 
Some infrared spectra of Protiwm Icicariba suggest the 
same relationship, whereas other species of Protiwm 
differ significantly. 


Colombian Amber 


We have examined amber from two localities in Co- 
lombia. Boussingault (1843) reported amber to have been 
found in large amounts in a gold-bearing alluvium at 
Giron, near Bucaramanga, Departamento de Santander 


[98 | 


del Norte. From the locality, this amber has been named 
Bucaramangite. Since it was found in alluvium, it might 
have been transported some distance from beds of any 
age. The sample analyzed came from a pale yellow trans- 
parent piece that weighed 12 kg. It is insoluble in alco- 
hol, swells and becomes opaque in ether and does not 
contain succinic acid. 

Four spectra were run from USN M collection #R7317. 
Three spectra (Plate X XVI) were similar to Type I 
(H 429). Another spectrum was like Type II (H 208) 
which had a better resolved version of Type I. 

Cockerell (1928) reports two species of Diptera in 
amber from the Valle de Jestis in the Departamento de 
Santander del Sur. He characterized it only as being 
‘‘relatively soft’’ and of ‘‘uncertain age. ”’ 

Amber from Medellin, Colombia, is also available from 
USNM collection #97466. Although it is listed as ‘‘co- 
palite,’’ we are unaware that its geologic age or its chemi- 
eal or physical description have been published. Six 
spectra were run and H 359 (Plate X X VI) is represent- 
ative of them. 

The spectra represented by H 208 (Girén) and H 359 
(Medellin) can be related generally to spectra of Hy- 
menaea Courbaril, just as in the case of amber from Chia- 
pas, Mexico. In previous discussions of Chiapas amber 
(Langenheim and Beck, 1965; Langenheim, 1966; 
Langenheim, 1967), spectra of resin from modern pop- 
ulations of Hymenaea Courbaril were compared from 
Mexico, Guatemala, British Guiana (Guyana), Brazil, 
Venezuela, and Ecuador. Considerable similarity in the 
spectra throughout the wide distribution of this popula- 
tion was noted, although certain populations were more 
similar than others. For example, spectra of the Chiapas 
amber resembled more closely the spectra of Hymenaea 
resin from populations in Mexico and Guatemala than 


[ 99 | 


those with a more southern distribution. Therefore, it 
was not surprising when the spectra of amber from Girén 
and Medellin, Colombia, compared closely with H. 
Courbaril resin from Guyana and Brazil (Plate XX VII). 
We presently do not have specimens of Hf. Courbaril 
from Colombia. 

These spectra of Colombian amber seem sufficiently 
close to present-day Hymenaea to indicate its possible 
botanic origin. Unfortunately, we do not know the geo- 
logic age nor have corroborative evidence for the botanic 
source from plants included in the amber or in the amber- 
bearing beds, such as was available for the amber from 
Chiapas, Mexico. 


[ 100 | 


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[ 105 ] 


[ 901 | 


WAVE NUMBER (CM-!) 
4000 3000 2000 1500 1000 900 800 700 


ALASKA | 


KREISCHERVILLE, NY. 


i | J J 


1 i i 


! 
6 7 8 9 10 1] 12 13 14 15 
WAVELENGTH (MICRONS) 


Infrared spectra of Cretaceous amber from Alaska Types I and II and Kreischerville, N.Y. 


el 
bh 
tht 


AIX Sivi1g 


[ LOL ] 


WAVE NUMBER (CM!) 


coo 99020001500 tp) so ago 


H 318 


ALASKA III 


WASHINGTON, D.C. 


GREENLAND 


J | 1 i ! J | ] 1 I 


L 
a 4 5 7 8 9 10 H 12 3. 14 +15 
WAVELENGTH — (MICRONS) 


Infrared spectra of Cretaceous amber from Alaska Type III, from Washington, D.C. and 


of unknown age from Greenland. 


aLVIg 


AX 


[ SOL | 


WAVE NUMBER (CM7!) 
4000 3000 2000 1500 1000 900 800 700 
L tL 


MANITOBA,CANADA AMBE 


AGATHIS AUSTRALIS RESIN H 265 


NEW ZEALAND AMBER 


1 | 


= 
Se 
= 
Lo 
& 


H 316 


3 4 


7 8 2 10 
WAVELENGTH (MICRONS) 


Infrared spectra of Cretaceous amber from Manitoba, Canada, Pleistocene amber from New 
Zealand, and of modern resin from Agathis australis. 


15 


ALVIG 


IAX 


[ 6OL ] 


WAVE NUMBER (CM-!) 
4000 3000 2000 1500 1000 900 
i] 


MAGOTHY RIVER, MD. H 36] 


VV 


KINCORA, NEW JERSEY 


CHARLESTON, S.C. 


! __ ! i } 


i ! i ! j 
3 4 5 6 7 8 9 10 1] 2 13 14 
WAVELENGTH (MICRONS) 


Infrared spectra of Cretaceous amber from Magothy River, Maryland; Kincora, N.J.; 


and Charleston, S.C. 


ALVT 


TIAN 


[ OLT | 


WAVE NUMBER (CM~') 


4000 3000 2000 1500 1000 900 800 700 


] j 


b— 


ST. GEORGES, DELAWARE H 357 


BORDENTOWN, N.J. 


H 422 
CLIFFWOOD, N.J. 


| \ 1 | i | 


H i | ! 


i 
4 2 6 7 8 9 10 11 12 3 14 15 
WAVELENGTH (MICRONS) 
Infrared spectra of Cretaceous amber from St. Georges, Delaware; Bordentown, N.J.; 


and Cliffwood, N.J. 


ALW Tf 


ITIAX 


{ LIL ] 


WAVE NUMBER (CM-!) 
2000 1500 1000 900 800 
! 1 


1 


ROEBLING, N. J. 


H 350 


H 420 


Wer 


i t | | L ! ! L L l 
5 


4 8 2 10 12 13 
WAVELENGTH (MICRONS) 


Infrared spectra or Cretaceous amber from Roebling, N.J. Types I, II and III. 


5 


ALV Ig 


XTX 


[ aI | 


= 


WAVE NUMBER (CM7!) < 

4000 3000 2000 1500 1000 900 800 700 > 
PEMBERTON N. J. ¥ 

i ne 


MONTANA | 


HARRISONVILLE, NJ. 


l 1 | _| 1 J i i J if 1 


4 > 6 7 8 - 10 1] 12 13 14 15 
WAVELENGTH — (MICRONS) 
Infrared spectra of Cretaceous amber from Pemberton, N.J. and Harrisonville, N.J. 
and unknown age from Montana Type I. 


[ SLL | 


WAVE NUMBER (CM-!) 


4000 3000 2000 1500 1000: 900 800 700 
l L 


! 1 I j 1 


ee 
MONTANA II So! 405 


al H 425 
WS , 
pe a a 


MARTHA’S VINEYARD. we NO m 


\ 


| i i i i L it i 


1 1 1 

4 2 6 F 4 8 9 10 12 13 14 15 
WAVELENGTH (MICRONS) 

Infrared spectra of amber of unknown age from Montana Types II and III 


and Cretaceous amber from Martha’s Vineyard, Mass. 


IXX avid 


[ FLL | 


WAVE NUMBER (CM~') 
4000 3000 2000 1500 1000 990 


BAJA CALIFORNIA, 
MEXICO 


S 
= 
= 
= 


i 1 J J ! I 


aR 
| III 


3 4 5 6 7 8 9 10 1] 12 13 
WAVELENGTH (MICRONS) 


Infrared spectra of Cretaceous amber from Baja, California, Mexico, Types I, II and III. 


TLW Td 


TIXX 


[ CLL J 


WAVE NUMBER (CM-!) 
4000 3000 2000 1500 1000 900 800 700 
l 


CHIAPAS, MEXICO eee 


7 a 


l BI 1 l L | | 1 ! 1 1 
4 5 6 7 8 9 10 1] 12 13 14 


WAVELENGTH (MICRONS) 


Infrared spectra of Oligo-Miocene amber from Chiapas, Mexico, Types I, II and III. 


15 


HWIXN &tv1d 


[ ott | 


\ 
= 


WAVE NUMBER (CM-1) 


4000 3000 2000 1500 1000 700 800 700 
A ee | 
SEATTLE, WASHINGTON | H 323 


SEATTLE, WASHINGTON II H A2A 
| 


Sa 
CANON DIABLO, ARIZ. =~ H 365 
| awe aan 


i ! 1! ! | ! 1 L 1 1 1 i 


2 4 5 6 rs 8 4 10 1] 12 13 14 15 
WAVELENGTH (MICRONS) 
Infrared spectra of Eocene amber from Seattle, Washington, Types I and II and amber 
of unknown age from Cafion Diablo, Arizona. 


ALVIg 


AIXNN 


[ LIL | 


WAVE NUMBER (CM7-!) 
4000 3000 2000 1500 1000 900 800 700 
l eae! | 


DOMINICAN REPUBLIC | 


DOMINICAN REPUBLIC II 


GUAYAQUIL, ECUADOR 


i 1 I I l i } ] J at ] 


3 4 5 6 7 8 4 10 im 12 13 14 15 
WAVELENGTH (MICRONS) 


Infrared spectra of Oligocene amber from the Dominican Republic, Types I and II and 
amber of unknown age from Guayaquil, Ecuador. 


ALV Ig 


AXX 


[ SEL | 


WAVE NUMBER (CM7!) 
1500 


ii 


| GIRON, COLOMBIA II 


Yew 


, H 359 
| MEDELLIN, COLOMBIA 


H 208 


J ! l ! L | 1 1 ! 1 ! tL 
3 4 2 6 7 8 , 10 ie 12 13 14 15 
WAVELENGTH (MICRONS) 


Infrared spectra of amber of unknown age from Girén, Colombia, Types I and II and from 


Medellin, Colombia. 


ALV Tg 


IAXX 


[ 61T | 


WAVE NUMBER (CM-!) 


4000 3000 2000 1500 1000 900 800 700 


i L tL 


CHIAPAS, MEXICO AMBER 


(MEXICO) By 


H.COURBARIL RESIN (GUYANA) 


H 58 


GIRON, COLOMBIA AMBE 


| | | ! | L ! 1 


= 


| 1 L 


3 4 5 6 7 8 9 10 1] 12 13 14 15 
WAVELENGTH (MICRONS) 


Infrared spectra of Oligo-Miocene amber from Chiapas, Mexico, modern resin from Hymenaea 
Courbaril from Mexico and Guyana and, amber of unknown age from Girén, Colombia. 


ALW Td 


WAX X 


OzI | 


[ 


4000 3000 2000 1500 
! L 


WAVE NUMBER (CM-1!) 
1000 900 800 700 
l 


| i 


HYMENAEA COURBARIL RESIN 
(COSTA RICA) 


PARA, BRAZIL AMBER 


H. COURBARIL RESIN (BRAZIL) 


J l I | | i rt i 


oP 


7 8 9 10 12 13 14 15 
WAVELENGTH (MICRONS) 


Infrared spectra of Miocene amber from Para, Brazil, and modern resin of Hymenaea Courbaril 
from Costa Rica and Brazil. 


ALVIg 


IIAXX 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


CamsBripce, Massacuuserrs, JANUARY 10, 1969 VoL. 22, No. 


DE PLANTIS TOXICARIIS E MUNDO 
NOVO TROPICALE COMMENTATIONES III 


PHYTOCHEMICAL EXAMINATION OF SPRUCE’S ORIGINAL 
COLLECTION OF BANISTERIOPSIS CAAPI 
BY 
Ricuarp Evans Scuuttes', Bo HoLMsreptr 
AND JAN-ErIK LINDGREN’ 


RicHARD Spruce, the humble Yorkshire schoolmaster- 
bryologist, became one of the outstanding tropical plant 
explorers of all time. On July 12, 1849, he arrived at 
the mouth of the Amazon to start his epoch-making 
botanical studies and collections that spanned a period of 
fifteen years (1849-1864) in the Amazon and the Andes. 
Spruce (Plate XXX) was far ahead of his day in 
scientific thought and method. He lived closely with 
native peoples, learned several languages and kept his 
mind ever inquisitive and his eye ever perceptive. For 
a number of plants that have later attracted extensive 
phytochemical and pharmacological attention — and 
which are still claiming serious studies —it was Spruce 
who gave us detailed, accurate, pioneer information. 
One of these plants was a jungle liana, source of an 
extraordinary hallucinogenic drink called caapz in Brazil, 
ayahuasca in Ecuador and Peru. It was in 1852, during 
' Botanical Museum of Harvard University, Cambridge, Mass. 


> Department of Toxicology, Swedish Medical Research Council, 
Karolinska Institutet, Stockholm, Sweden. 


[ 121 ] 


the early stages of his five years of field work on the upper 
Rio Negro in Amazonian Brazil, that Spruce first learned 
of caapi amongst the Tukanoan tribes of the region. It 
was employed to induce, for prophetic and divinatory 
purposes, an intoxication characterized, amongst other 
strange syndromes, by frighteningly realistic colored 
visual hallucinations and a feeling of extreme and reck- 
less bravery. Unlike many early reports of newly dis- 
covered narcotics, Spruce’s contribution included a 
precise determination of the botanical source of the drug. 

Finding caapi cultivated along the Rio Negro, he 
noted that ‘“‘there were about a dozen well growing 
plants... twining up to the tree tops... and several 
smaller ones. It was fortunately in flower and young 
fruit; and I saw, not without surprise, that it belonged 
to the ... Malpighiaceae ...*’ A collection in full 
flower (Spruce 2712) was taken from the liana, and he 
drew up a description of the species from living speci- 
mens. He allocated the species to the genus Banisteria, 
calling it Banisteria Caapt from the vernacular name. 
This description was published by the botanist Grisebach. 
As taxonomic understanding of the family grew in the 
present century, the American specialist, C. V. Morton, 
ascertained that this species-concept could not with pre- 
cision be included in Banisteria, and, in 1931, he trans- 
ferred it to the genus Banisteriopsis. The liana is, accord- 
ingly, now correctly called Banisteriopsis Caapi (Spruce 
ex Grisebach) Morton. * 

Even a century ago, Spruce’s thinking was, at least in 
part, along chemotaxonomical directions. He mused: 


*Banisteriopsis Caapi (Spr. ev Griseb.) Morton in 
Journ. Wash. Acad. Sci. 21 (1931) 485. 

Banisteria Caapt Spruce ex Grisebach in Martius FI. 

Bras. 12, pt. 1 (1858) 43. 


[ 122 ] 


Plate XXIX 


BANISTERIOPSIS Caapi 
( Spruce ex Griseb,) Morton 


7; 


iy 7. 
Wii Pon, "Ys \ 


1, flowering branch, about $ . 2, flower, about 24 *. 38, fruit, 
somewhat larger than $ ™., 


‘*My surprise arose from the fact that there was no nar- 
cotic malpighia on record, nor indeed any species of that 
order with strong medicinal properties of any kind. 
Byrsonima ... includes many species... their bark 
abounds in tannins... Another genus—Bunchosia .. . 
of the Andes. . . is described in books as poisonous, and 
if it be really so, then it is the only instance, so far as | 
know, of the existence of any hurtful principle in the 
entire family ... excepting ... caapi.”’ ‘‘Yet,’’ he 
prophetically remarked ‘‘strong poisons may lurk undis- 
covered in many others of the order, which is very large 

; and the closely allied soapworts (Sapindaceae) 
contain strong narcotic poisons, especially in the genus 
Paullinia.”’ 

In many ways, Spruce was ahead of his times. In 
those years, there was little liaison between botanical 
explorers and chemists of the laboratory. Botanists sel- 
dom gathered material for phytochemical study, and in 
Spruce’s case the great distance and isolation of his scene 
of field work and the primitiveness and absence of nor- 
mal communications one might believe would have made 
it impracticable or impossible for him to gather material 
in bulk for pharmaceutical specialists. Notwithstanding 
these drawbacks, Spruce did so enrich science, but, like 
so many collectors even in modern times, he was frus- 
trated in his attempt. 

‘*T obtained a good many pieces of stem [from the 
type plant of Banisteria Caapi], dried them carefully, 
and packed them in a large box, which contained the 
botanical [herbarium] specimens, and dispatched them 
down the river for England in March 1853. The man 
who took that box and four others on freight, in a large 
new boat he had built on the Uaupés, was seized for debt 
when about half-way down the Rio Negro, and his boat 
and all its contents confiscated. My boxes were thrown 


[124 J 


PLATE XXX 


RICHARD SPRUCE 


Drawn by Kimer W. Suivi 


aside in a hut, with only the damp earth for floor, and 
remained there many months, when my triend Senhor 
Henrique Antoni of Manos... succeeded in redeem- 
ing them and getting them sent to the port of Para. 
When Mr. Bentham came to open them in England, 
he found the contents somewhat injured by damp and 
mould, and the sheets of specimens near the bottom of 
the boxes quite ruined. The bundle of the caapi would 
presumably have quite lost its virtue from the same 
‘cause, and I do not know that it was ever analyzed 
chemically ; but some portion of it should be in the Kew 
Museum at this day.” 

In an address at the [IT International Pharmacologi- 
‘al Congress in Sio Paulo, Brazil, in 1966, one of the 
authors (2, 2a) said: *‘One of the most interesting exer- 
cises that I can imagine would be the analysis of a small 
portion of this original Spruce material—if, indeed, it is 
still preserved at) Kew—with our modern improved 
chemical techniques. The active principles of caapi (har- 
mine type alkaloids) might not have deteriorated with 
the mildew, and it is possible that even in this more than 
a century of storage, the alkaloids would be intact.” 

In later discussions of this interesting experiment, we 
resolved to try to follow it up. Accordingly, on Decem- 
ber 21, 1967, we wrote to Sir George Taylor, Director 
of the Royal Botanic Gardens, Kew, and requested a 
small amount of the stem material of Banisteriopsis Caapi 
to which Spruce had referred, if it still existed. Shortly 
thereafter, Sir George and Dr. Patrick Brenan, Keeper, 
located this valuable historical collection and, on April 
26, 1968, sent a letter informing us that some of this 
material would be made available for chemical study. 
Also enclosed were notes on the material from the entry 
book for 1854. The label on the specimens states: 

“Stems of Banisteria sp. used with the roots and leaves 


[ 126 ] 


of Haemadictyon in the preparation of an intoxicating 
beverage called ‘Caapi’. Rio Uaupes. PR. Spruce 166. 
HE 2712.” 

The entry in the notebook reads as follows: 

**Portions of the stems of a Malpighiaceous twiner, 
apparently an undescribed Banisteria (2712 to Benth.), 
‘alled by the Indians Caa-pi: and of the roots and leaves 
of a Haemadictyon, called Caapi-pinima (i.e. ‘painted 
‘aapi ) the leaves being veined with red. From these in- 
gredients, the Banisteria entering much more largely 
than the Haemadictyon, is prepared an intoxicating 
drink known to all the natives on the Uaupés by the 
name of Caapi. 

“In the Dabocurés (or festas) of the Uaupé Indians, 
the young men who figure in the dances drink of the 
Caapi 5 or 6 times during the night, the dose being a 
cuya, the size of a very small teacup, twice filled. In 
two minutes after drinking it, its effects begin to be ap- 
parent. The Indian turns deadly pale, trembles in every 
limb, and horror is in his aspect: suddenly contrary 
symptoms succeed—he bursts into a perspiration and 
seems popeyed with reckless fury—seizes whatever arms 
are at hand, his murucu, cutlass, or bow and arrows, and 
rushes to the doorway, where he inflicts deadly wounds 
on the ground or doorposts, calling out ‘Thus would | 
do to such a one (naming some one against whom he has 
a grudge) were he within my reach.” In the space of 10 
minutes, the effect passes off, and the Indian becomes 
‘alm, but appears much exhausted. ”” 

The parts of Spruce’s material that reached the De- 
partment of ‘Toxicology, Karolinska Institutet, Stock- 
holm, in April 1968 consisted of five pieces weighing in 
all 26.7 g. (Plate NN NI); 11.5 g. were worked up for 
analysis by gas chromatography-mass spectrometry and 
other methods as described earlier (8, 4). The yield of 


L127 | 


Banisteriopsis Caapi (Spr. ex Griseb.) Morton. BR. Spruce No. 166. 


LLW'T 


ix 
NA LZ 


alkaloids was found to be 0.4 percent. A newly collected 
botanically verified specimen of Banvzsteriopsis Caapt 
analyzed at the same time was found to contain 0.5 per 
cent alkaloids (5). The latter material contained as de- 
scribed by many authors the main alkaloids harmine, 
harmaline and tetrahydroharmine (6). In addition, it 
contained two minor components that will be described 
separately (5). By contrast, the alkaloid content of the 
Spruce material consisted exclusively of harmine. This 
was proven beyond any doubt by gas chromatography 
and the combination of gas chromatography-mass spec- 
trometry. As evident from both the gas chromatograms 
(Plate X XXII) and the mass spectra (Plate XX XIII), 
there is complete identity between synthetic harmine 
and the alkaloid in Spruce’s material. It is open to 
question whether the stems sent home by Spruce in 
1858 from the beginning contained only harmine or 
perhaps more likely that harmaline and tetrahydrohar- 
mine have with time been transformed into the chemi- 
cally more stable aromatic $-carboline, harmine. 

Under any circumstances, it is remarkable that 
Spruce’s query about the chemical analysis of the ma- 
terial that fared so badly on its way from the Amazonian 
rain forest to the Royal Botanic Gardens at Kew has 
been answered by modern analytical microtechniques 
115 years later. 

This investigation was supported by grant MH 12007- 
03 (Holmstedt) and by grant LM-GM _ 00071-01 
(Schultes) from the National Institute of Mental 
Health, U.S. Public Health Service. We are indebted 
to Sir George Taylor, Director of the Royal Botanic 
Gardens, Kew, Surrey, England, for supplying us with 
the material from Spruce’s original collection. 


[ 129 ) 


PLaTE XXXII 


| BANISTERIA SP 
R.SPRUCE Nol66 


0 10 20 
MINUTES 


HARMINE 


|| OGG: 


cH, 


| | | 
| pA 
ee | ey ee a 


oO 10 20 
MINUTES 


Gas chromatogram of alkaloid fraction from Spruce’s material and 
reference substance. Conditions: Column 2 m ; i.d. 8.2 mm ; 59% 
y ry x te) 
OV-17 on 100-120 mesh Gas Chrom P ; temp. 230°. 


[ 130 | 


PuaTE XXXIII 


90 + | BANISTERIA SP 
R SPRUCE No. 166 


Relative intensity % 
w S.] 
[e) ° 


5.8 


———________, 


HARMINE 


8 
— 
| 

= 


8 8 


8 8 
T T 


1S.) 
oO 


> 
{e) 


Relative intensity % 
8 6 


3 


50 ere) 150 200 


Mass spectra of peaks shown in Plate XXXII. Conditions as 
described in Plate XXXII. 


[ 131 ] 


REFERENCES 


. Spruce, R. 4 botanist on the Andes and Amazon. |Ed. A.R. Wal- 


lace| Macmillan and Co., Ltd., London, 2 (1908) 414-430. 


. Schultes, R.E. ‘‘Some impacts of Spruce’s Amazon explorations 


on modern phytochemical research.’’ Rhodora 70 (1968) 313-339. 


7 i 66, : 4 ; 
.Schultes, R.E. “The impact of Spruce’s Amazon explorations on 


modern phytochemical research.’’ Ciencia e Cultura 20 (1968) 87- 
49. 


Holmstedt, B. and J.E. Lindgren in Ethnopharmacologic search for 
psychoactive drugs. {Ed. D. Efron] U.S. Public Health Service 
Publ. No. 1645 (1967) 339-373. 


Agurell, S., B. Holmstedt, J.E. Lindgren and R.E. Schultes. 
Acta Chemica Scandinavica. In press. 


Agurell, S. and J.E. Lindgren. To be published. 
Deulofeu, V. in Ethnopharmacologic search for psychoactive drugs. 


[Ed. D. Efron] U.S. Public Health Service Publ. No. 1645 (1967) 
393-402, 


{ 182 | 


DE PLANTIS TOXICARIIS E MUNDO 
NOVO TROPICALE COMMENTATIONES IV 
BY 
RicHarp Evans SCHULTES 


IN CONTINUATION of a series of articles on toxic plants 
of the New World tropics, this contribution will call 
attention to a number of species with miscellaneous in- 
teresting ethnobotanical notes as to their use but which, 
for the most part, have not been subjected to phyto- 
chemical examination. 

Most of the data reported in the following pages were 
gathered during field work which I have carried out in 
the American tropics or which my students have reported 
as a result of their ethnobotanical studies in sundry parts 
of Latin America. 

It will be clear to phytochemists that the analysis of 
some of the species enumerated may be of special inter- 
est, since a number of the plants belong to genera or 
even families from which biodynamic constituents have 
not hitherto been isolated. 

The current phases of this research on toxic plants of 
the New World tropics, including the preparation and 
publication of most of the plates in this contribution, 
are supported by a grant from the National Institutes of 
Health (No. LM-GB 00071-01). The line drawings have 
been prepared by the late Mr. John Stanwell-Fletcher 
and by Mr. Gordon W. Dillon, Mr. Joshua B. Clark and 
Miss Irene Brady. 

[ 133 ] 


The enumeration of families follows the Engler-Prant] 
system. 
GARRYACEAE 


Garrya laurifolia (Hartw.) Bentham var. macro- 
phylla (Hartw.) Wangerin in Pflanzenr. 4, Fam. 56a 
(1910) 16. 

Mexico: Estado de Oaxaca, Cerro Zempoaltepetl, eastern slope. 


Alt. 2600 m. ‘‘Very large shrub along forest trails.’ May 26, 1939. 
R.E. Schultes 547. 


The Mie Indians living near the base of Cerro Zem- 
poaltepet] employ the leaves and stems in the form of a 
decoction as a febrifuge, but when taken in excess this 
medicine reputedly causes a burning sensation in the di- 
gestive tract and a strong trembling of the hands. 

Garrya laurifolia, known in Mexico as cuauhchichic, 
is employed therapeutically in tincture or extract form 
in the treatment of chronic diarrhoea. Several analyses 
have been made, and it has been established that the 
bark infusion kills rabbits by paralysis of the respiratory 
centres(Martinez, M. : ‘‘Plantas medicinales de Mexico” 
ed. 4 (1959) 98-96). Amongst other constituents, it is 
said to contain a bitter alkaloid principle, garryine, a 
resin, tannic acid and a compound possibly of glycosidal 
nature. Several alkaloids have been isolated from the 
genus Garrya, including G. laurifolia var. macrophylla 
(Heérissey, H. and C. Lebas in Journ. Pharm. Chim. 2 
(1910) 490; Oneto, J.F. in Journ. Am. Pharm. Assoc. 
35 (1946) 204). 

ANNONACEAE 


Unonopsis veneficiorum (Mart.) R.E. Fries in 
Acta Hort. Berg. 12 (19387) 238. 
CotomsBia: Comisaria del Putumayo, Rio Guamiies, Santa Rosa. 


‘‘Primary forest. Small tree more or less 20 ft. Bark of root is used 
in preparation of curare.*” November 26, 1966. H.V’. Pinkley 558. 


[ 134 ] 


PuLaTE XXXIV 
UNONOPSIS venificiorum (Mart) RE.Fr. 


. * . 9 * 
1, habit, approximately + <. 2, flower, approximately 25 . $8, immature 


fruits, approximately 25 . 4 and 5, seed, approximately natural size. 6, 
fruits, approximately 4+ ». 


Mr. Homer V. Pinkley, one of my graduate students 
who spent more than a year in ethnobotanical studies 
amongst the Kofin Indians of Kcuador and Colombia, 
reports that the root of this small tree, known in Kofin 
as i-te-si-fan-di, is the source of a curare preparation of 
the tribe. 

This species was first reported as an arrow poison in- 
gredient by Martius (Spix, J.B. and K.F.D. Martius 
‘*Reise in Brasilien’ (1881) 1287), who stated that the 
Juri, Mirana and other Indians of the Rio Japura (Rio 
Caqueti) and Rio Negro of Colombia and Brazil so em- 
ployed it. Unonopsis veneficiorum has been reported as 
containing bisbenzylisoquinoline alkaloids (Hegnauer, 
R. “*Chemotaxonomie der Pflanzen’? 8 (1964) 118). 

Mr. James W. Walker of the Gray Herbarium of 
Harvard University has identified the collection Pinkley 
558. 


CONNARACEAE 


The chemistry of the Connaraceae is very poorly 
known and most certainly represents one of the areas in 
the angiosperms where phytochemists should concen- 
trate attention. Dye materials and balsamic resins are 
rather common in the family, and, although very poison- 
ous substances have been reported from the fruit, seeds 
and bark of some species, their chemical nature is not 
yet clarified (Hegnauer, loc. cit.) 3 (1964) 545-546. As 
a family very closely allied to the Leguminosae, the 
Connaraceae should be expected to possess 2 good num- 
ber of species employed as poisons in primitive societies. 

My identification of the connaraceous species men- 
tioned in this paper has been verified by Dr. Gilian T. 
Prance of the New York Botanical Garden. 


Connarus opacus Schellenberg in Engler Pflanzenr. 
4, Fam. 127 (1988) 244. 


[ 136 | 


PLATE XX AV 


A 
Fan 
A i 
i ty /\ “i 
/ : 
a 
/ ~ 
\ Al 
hs } noe -\ 
f Jes 
ih ayy “| TN 


Pe 
CONNARUS 4 
OPrucel SRO 
: LBaker >\\ 


ane 
= 


i i 
i 


’ 
“a ft 
ih) 


CONNARUS 


opaec us 


Habit, approximately 4 X. 


Cotomsia: Comisaria del Vaupés, Rio Kuduyari, headwaters. **Li- 
ana. Fruits dull red. Bark and leaves used by Kubeo for fish poison.’’ 
August 16, 1960. R.E. Schultes 22719, 

This extensive vine, which is relatively abundant in the 
headwaters of the Rio Kuduyari as an element of the 
riverside vegetation, provides one of the usual fish poi- 
sons employed in still water by the Kubeo Indians. The 
root and stem bark and leaves are mixed together, 
crushed, placed in a bag and drawn slowly through the 
water. 

A spot test for alkaloids, made on the dried leaves of 
this collection, was negative. Alkaloids have apparently 
never been reported from any species of the Connaraceae 
(Willaman, J.J. and B.G. Schubert: ‘‘Alkaloid-bearing 
plants and their contained alkaloids’? Techn. Bull. No. 
1234, U.S.D.A. (1961). 

The collection Schultes 22719 is apparently the first 
record of this species from Colombia. The type was col- 
lected in British Guiana by Schomburgk. 


Connarus Schultesii Standley ex Schultes in Bot. 
Mus. Leafl., Harvard Univ. 9 (1941) 173, t. 4. 

Mexico: Estado de Oaxaca, San Juan Lalana, Distrito de Choapam. 
Alt. 700 m. ‘SA tangled vine growing on large forest trees, mountain 
southeast of Lalana, Stems exude red sap upon breaking.’’ May 9, 
1939. R.E. Schultes et B.P. Reko 833. 

Known only from the type collection, this species has 
been employed by the Chinantec Indians as a taenifuge. 
The extremely bitter red sap that exudes from the stems 
acts in the expulsion of certain intestinal parasites, but 
it is reputedly toxic in large doses. Since the species is 
reported by the natives to be local, its use in folk-medicine 
appears to be restricted. It is perhaps of interest that an 
African species is similarly employed (Heckel, E., and 
I. Schlagenhauffen in Ann. Fac. Sci. Marseilles 6, fase. 
2 (1897) 1-26). The only other species recorded from 


[ 188 ] 


PuaTE XXXVI 


CONNARUS 


Schultesii 


Standley 


enlarged approximately 14 X. 


1, fruiting branch, approximately 4 . 2, fruit, 


Mexico—Connarus lentiginosus Brandg. of Chiapas—is 
not known to be utilized medicinally, but ethnobotanical 
studies have not been pursued in the region. 

The red sap of Connarus Schultesii may contain color- 
ing constituents, but the Chinantecs and neighboring 
Zapotecs apparently do not make use of them as a dye. 


Connarus Sprucei Baker in Martius FI. Bras. 14, 
pt. 2 (1871) 187. 
Cotomsia : Comisaria del Vaupés, Rio Kuduyari, near mouth. ‘‘Vine. 


Flowers white. Bark employed as fish poison.’’ August 12, 1960. 
RE. Schultes 22576. 


The bark of the root and stem of this riverside liana 
is employed by the Kubeo Indians as a fish poison. 

Spot tests for alkaloids, made on the bark of the spe- 
cies, proved to be negative. 


Rourea glabra Humboldt, Bonpland et Kunth. Nov. 
Gen. et Sp. 7 (1825) 41. 
CoLomsia: Comisaria del Amazonas, Rio Karaparana, between the 


mouth and El] Encanto. Alt. about 150 m. ‘‘Small tree. Fruit red 
with blackish blue husk.’? May 22-28, 1942. R.E. Schultes 8831. 


The Witoto Indians use the root and stem bark of 
Rourea glabra as a fish poison. This small tree or vine is 
rather common in pockets along the flood-banks of the 
Rio Karaparana. Although it is a widespread species, 
occurring in Central America, the West Indies and tropi- 
cal South America, no uses seem to have been reported 
for it beyond the employment of a decoction as a gargle in 
treating catarrhal conditions (Schellenberg, loc. cit. 214). 
Two vernacular names for the plant in Brazil—mata ca- 
chorro (“dog killer’’) and mata negro (‘‘negro killer’*)— 
do, however, suggest poisonous properties. The fruit and 
root bark of Rourea erecta (Blanco) Merr., an Asiatic 
species, are reputedly poisonous to dogs and other car- 


| 140 | 


PiuateE XXXVII 


ROUREA gl 


\! 
\" 


abra HBK. 


ae 
Habit, approximately 4 X. 


nivorous animals (Brill, H.C. and A.H. Well in Philip- 
pine Journ. Sci. 12A (1917) 171). A steroidal sapogenine 
has been found in the leaves and twigs of Rourea ligu- 
lata Bak. of Brazil (Altman, R.F.A. in Nature 178 
(1954) 1098). 

MUPHORBIACEAE 


Phyllanthus lathyroides Humboldt, Bonpland et 
Kunth Nov. Gen. et Sp. 2 (1817) 110. 


Mexico: Estado de Oaxaca, Distrito de Teotitlan, Huautla de 
Jiménez. August 3, 1938. R.F. Schultes et B.P. Reko 417. 


This common roadside weed, called shha-nin-du 
(‘‘eruption plant”’) in the Mazatee Indian language, has 
medicinal properties, according to the natives. A decoc- 
tion of the leaves is used as a wash for eye infections, 
such as conjunctivitis. A poultice of moistened leaves is 
applied to boils, hence the Indian name of the plant. A 
strongly emetic tea is, likewise, prepared from the leaves. 


FLACOURTIACEAE 


Lunania parviflora Spruce ex Bentham in Journ. 
Linn. Soc. 5, Suppl. 2 (1861) 90. 


Co.tomsBia: Comisaria del Amazonas, Trapecio Amazoénico, Rio Lore- 
’ I 
toyacu. September 1946. R.E. Schultes et G.A. Black 8316. 


The Tikuna Indians report that the root of Lunania 
parviflora is toxic and was formerly powdered and added 
to cooked food for the purpose of killing enemies or un- 
wanted visitors from foreign tribes. 


THYMELAEACEAE 
Schoenobiblus peruvianus Standley in Field 
Mus. Publ. Bot. 11 (1986) 169. 


Cotompia: Comisaria del Putumayo, Rio Sucumbios (San Miguel), 
Conejo y alrededores, frente a la Quebrada Conejo. *“Large shrub. 
Root furnishes one of the poisons for curare.’*’ April 2-5, 1942. R.E. 


[ 142 ] 


PLaTE XXXVIII 


SCHOENOBIBLUS _ peruviagnus -— Standi. 


of SOM, 
S 


QA NNN 
We AY 
rr 


1, habit, approximately 2 <. 2, leaf and bractlet, approximately 2 <. 3, 
flower, approximately 4 . 4, flower, approximately 6 X. 5, fruit, approxi- 
mately 5 X. 


Schultes 3521. —** Poison. Fruits used in making curare. Shrub.” April 
2-5, 1942. Schultes 3655,.—Santa Rosa y alrededores. *‘Roots and 
fruits used in Kofan curare preparation.’’ April 7-8, 1942. Schultes 
3613,—Rio Guamiies, Santa Rosa. Alt. 990 ft. ‘“Low and swampy 
area. Plant 44 ft. tall. Fruit bright orange. Root and fruit used in 
arrow poison; also sometimes used as a fish poison.’’ September 6, 
1966. H.V. Pinkley 435. 

In 1949, Schultes (in Bot. Mus. Leafl. Harvard Univ. 
13 (1949) 285-289) reported the use of Schoenobiblus 
peruvianus as ‘‘a very important ingredient in the arrow- 
poisons of the Kofin Indians of the Putumayo’’ in Co- 
lombia along the Ecuadorian border. The roots and the 
fruits are employed. 

Mr. Pinkley, asa result of his year-long ethnobotanical 
studies amongst the Kofiin Indians, has been able to 
conclude a detailed study of the employment of this in- 
teresting toxic plant in the preparation of arrow poisons, 
areport of which will be published elsewhere. His record 
that the plant is sometimes utilized as a fish poison 
extends our knowledge of its toxic activity and may be 
of interest chemically, inasmuch as the identity of the 
active principle is, as yet, unknown. 

The identification of the specimens cited above has 
been checked by Dr. Loren I. Nevling of the Arnold 
Arboretum. 

STYRACACEAE 


Styrax Tessmannii Perkins in Notizbl. 10 (1928) 
459. 


Cotombia ; Comisaria del Amazonas, Trapecio Amazonico, Rio Lore- 
toyacu. Tikuna name: me-re-ta-kee. March 1946. R.E. Schultes 7144. 


According to Tikuna Indians living along the Rio 
Loretoyacu, the aromatic resin from the stems of Styrax 
Tessmanni is poisonous when ingested. This has appar- 
ently not hitherto been reported, but, in view of persist- 
ent information to this effect, it bears phytochemical 
investigation. 

[ 144 ] 


PLaTteE XXXIX 


Flercher. 


6 


STYRAX 


Tessmannii Perk. 


1, habit of flowering branch, approximately 4 2, habit of fruiting branch, 
ot 


x. 
approximately $<. 3, flower, approximately 24 X. 4, fruit, approximately 4 x, 


GENTIANACEAE 


Chelonanthus alatus (4ub/.) Pulle Enum. Vase. 
Pl. Surinam (1906) 876. 


Mexico: Estado de Oaxaca, Distrito de Cuicatlan, between San 
Juan Zautla and Teutila. June 30, 1989. R. FE. Schultes 731.— Distrito 
de Choapam, San Juan Lalana, May 8, 1939. R.E. Schultes et B.P. 
Reko 7805. 

Cotombia: Comisaria del Amazonas, Rio Karaparand, between the 
mouth and El Encanto. May 22-28, 1942. R.E. Schultes 3805.— 
Comisaria del Vaupés, Rio Vaupes, Mitiand vicinity. September 27- 
October 20, 1966, R.E. Schultes, R.F. Raffauf et D. Soejarto 24246a 
Schultes, Raffauf et Soejarto 24346, 

Peru: Departamento de Loreto, Rio Napo, Negro Ureo. August 
16, 1966. R.T. Martin et C.A. Lau-Cam 1286. 
1290. 


Martin et Lau-Cam 


The Chinantecs in the Districts of Choapam, Ixtlin 
and Cuicatlin, Oaxaca, state that the root of Chelonan- 
thus alatus is bitter and is used in decoction as a febri- 
fuge, for cramps and indigestion, and to treat sores in 
the mouth. It has uses similar to those of Lisianthus 
migzrescens reported below. Applications based on its 
bitter principles are recorded from French Guiana 
(Heckel, KE. ‘‘Les plantes médicinales et toxiques de la 
Guyane Frangaise’* (1897) 57). Aublet, who described 
the concept, reported its employment in folk medicine 
in French Guiana as early as 1775 (Aublet, J.B.C.F. 
‘*Histoire des plantes de la Guiane Francaise’ 1 (1775) 
205), stating: ‘‘The whole plant is bitter. It is used to 
treat obstructions, and | have employed it for this pur- 
pose with success”’. 

The Witoto Indians of Amazonian Colombia, who 
refer to the plant as ho-/0-so-gi-né, dry and pulverize the 
leaves and flowers for powdering clothing and bedding 
to ward off insects. 

Kield spot tests on fresh material in Colombia have 
indicated that no alkaloids are present in the plant. 


[ 146 ] 


PuaTE XL 


— os _ 
_ CHELONANTHUS 
A alatus | 


(Aubl) Pulle- 


Habit, approximately 4+ . Flower dissected, approximately natural size. 
Anther, approximately 64 . 


Several specimens collected in eastern Peru (Martin 
et Lau-Cam 1286 and 1290) reported a vernacular name 
amaraguna which presumably refers to the bitter prin- 
ciples in this plant. Another name employed for this 
species in eastern Peru is una de tigre. A Peruvian col- 
lection (Mewia 4153), referable to Chelonanthus chelo- 
noides (.) Gilg, records that the plant is ‘‘used as a 
remedy for worm-infested wounds in cattle’’. 


Lisianthus nigrescens Chamisso et Schlechtendahl 
in Linnaea 9 (1831) 888. 


Mexico: Estado de Oaxaca, Distrito de Choapam, Santiago Zaca- 
tepec. R.E. Schultes 514.—Between Choapam and San Juan Comal- 
tepec. June 3, 1939. Schultes 573.—Tonanguilla. June 4, 1939. 
Schultes 591.—Estado de Oaxaca, Distrito de Choapam, San Juan 
Lalana. May 6, 19389. R.E. Schultes et B.P. Reko 772.—Between 
Latani and Choapam. May 13, 1939. Schultes et Reko S89. 


This striking, almost black-flowered herb occurs widely 
in northeastern Oaxaca, where it enjoys a variety of folk 
uses. Amongst the western Chinantees, a decoction of 
the roots is employed to relieve indigestion and _ heart- 
burn and as a febrifugal tonic. The neighbouring Maza- 
tecs apparently utilize the plant medicinally only ‘‘contra 
el aire’. The Mie Indians of Zacatepec apply a poultice 
of the leaves in the treatment of fungal infections of the 
skin of the feet, ankles and hands and value a decoction 
of the root asa bitter and febrifuge. Amongst the Zapo- 
tecs of Villa Alta and vicinity, the black, sticky flowers 
of this species are prepared in a wash employed to remove 
pinolillos (ticks) from the body, a use that is the basis of 
the Zapotec names le-zia and o-zia (‘‘remedy against 
ticks”’). 

Lisianthus nigrescens has apparently not hitherto been 
reported as auseful plant, although other species, known 
to possess the bitter properties characteristic of the 
Gentianaceae, find medicinal employment in various 


[ 148 ] 


, LISIANTHUS 


or | 


iI nigrescens 


Cham. ¢ Schlechtd. 


i) lo 
o wila 


Habit, approximately 4 ><. Flower dissected, approximately natural size. 
Anthers, approximately 10 *. 


[ 149 | 


parts of tropical America. The Brazilian Liseanthus pen- 
dulus Mart., for example, is valued asa bitter tonic and 
febrifuge; several other species are similarly used in 
French Guiana (Heckel, loc. cit., 57). The insectifugal 
properties of Listanthus nigrescens seem not to have been 
recorded, and, in view of a similar use of a species of the 
closely allied genus Chelonanthus (see above), they would 
appear to be worth investigating. 

In northeastern Oaxaca, this plant is known amongst 
the Chinantecs as /ee-the. ‘The Mazatecs of Huautla refer 
to it as shka-tee-tso. The Mije call it yerh. 


GESNERIACEAE 


Nautilocalyx sp. 


Cotoms1a: Comisaria del Vaupés, Rio Vaupés, savannah at base of 
Cerro de Mitt. September 27—October 20, 1966. R.E.Schultes, R.F. 
Raffauf et D. Soejarto 24211. ‘‘Flowers white. Terrestrial.’’ 


A spot test for alkaloids, given on fresh material in 
the field, was positive. This result is noteworthy in view 
of the fact that evidence of alkaloids in this family is 
almost wholly negative. An unknown alkaloid has been 
reported from the European Ramondia pyreaica Rich. 
Alkaloid spot tests made on several species of Besleria 
(B. ignea Fritsch: Schultes, Raffauf et Soejarto 24053) 
and Alloplectus (A. semicordatus P. et E.: Schultes, 
Raffauf et Soejarto 24225) from the Colombian Amazon 
were negative. 

The collection Schultes, Raffauf et Soejarto 24211 is 
sterile. It has been identified to genus by Dr. H. Emery 
Moore of the Bailey Hortorium. 


RUBIACEAE 


Duroia L. fil. 
Although this genus has apparently not been recorded 
as toxic, the number of reports from natives affirming 


[ 151 | 


the poisonous properties of the seeds of several species, 
all reports from widely separated localities, calls attention 
once again to the need for phytochemical studies guided 
by ethnobotanical observations. 


Duroia hirsuta (P. et E.) K. Schumann in Martius 
KF]. Bras. 6, pt. 6 (1889) 367. 
Cotompia: Departamento del Cauca, Puerto Limon, bosques en el 


lado caucano del Rio Caqueta’ Alt. 300 m. “‘Small tree. Bark, tied 
on arm, forms blisters, Soliman.’’ February 28-29, 1942. R.E. Schultes 


3320,—Comisaria del Putumayo, Rio Sucumbios, Conejo and vicinity. 
** Kofan = sha-ka-ker-nd-se’? April 2-5, 1942.—Comisaria del Ama- 
zonas, Rio Caucaya, between Puerto Jaramillo and Rio Putumayo. 
May 16, 1942. R.E, Schultes 3712.—Comisaria del Vaupés, Rio Apa- 
poris, near confluence of Ajajuand Macaya. Soliman. *‘January 1944. 
G. Gutiérrez et R.E. Schultes 612.—Intendencia del Meta, Sabanas 
de San Juan de Arama, margen izquierda del Rio Giejar, alrededores 
de aterrizaje Los Micos.’’ Alt. about 500 m. “‘Arbolito de 3 m. de 
alto. Comun en bosque.’’ December 5-20, 1950. J.M. Idrobo et R.E. 
Schultes 594, 


The type collection of Duroia hirsuta was made in 
Villavicencio, Colombia. Idrobo et Schultes 594, conse- 
quently, from near the town of Villavicencio, may be 
considered topotypical. 

Amongst many Indians of the Colombian Amazon 
and Putumayo—especially the Kofiin, Siona, Witoto 
and ‘Tikuna—the caustic bark of this small tree is em- 
ployed to make bluish black markings on arms and legs. 
Strips of the freshly removed, pliable bark are tied onto 
the arm or leg with the inner surface of the bark touch- 
ing the skin. After several hours or half a day, it is re- 
moved. Blisters and a localized red swelling result in 
another two to three hours. When this condition dis- 
appears, a dark band persists and lasts for several weeks, 
even occasionally for several months. There have appar- 
ently been no chemical analyses directed towards a dis- 
covery of the caustic principle in Duroia hirsuta. 


[ 152 ] 


PLateE XLII 


¥ 
Won 
WW Rj 


DUROIA _ hirsuta 
\ (P&E) Schum. 


1 and 2, habit, approximately 4 . 3, flower, approximately 1} . 


This small tree, known throughout eastern Colombia 
as soliman, is believed by the Indians to ‘‘poison”’ other 
plants. It is found usually in colonies of up to twenty 
individuals in the forest, and invariably nothing grows 
underneath the tree, except perhaps Selaginella (see 
photograph, Plate XN VIII, lower figure, in Bot. Mus. 
Leafl., Harvard Univ. 15 (1951) ). The natives explain 
this curious ecological phenomenon of the absence of 
vegetation by saying that the roots of Duroita hirsuta 
‘“*poison’’ the other plants. The real reason may be con- 
nected with the presence in Duroia hirsuta of swollen 
internodes which are always inhabited by ants. 


Duroia kotchubaeoides Steyermark in Mem. N.Y. 
Bot. Gard. 12 (1965) 201. 

Co.ompia: Comisaria del Vaupés, Rio Guainia, Puerto Colombia 
(opposite Venezuelan town of Maroa) and vicinity. Alt. about 800- 
850 ft. “"Small tree. Flowers:white.*’ October 31—November 2, 1952. 
R.E. Schultes, R.E.D. Baker et I. Cabrera 18211. 

Amongst the inhabitants of the Rio Guainia, the seeds 
of Duroia hotchubacoides are held to be poisonous when 
saten. 

Schultes, Baker et Cabrera 18211 is the type collection 
of this curious species. It is known only from one other 
collection made from the upper Orinoco in Venezuela. 
The species appears to be a restricted endemic. 


Duroia petiolaris (Spr. ) Hooker fil. ex K.Schumann 
in Martius Fl. Bras. 6, pt. 6 (1889) 364. 

CoLomaia : Comisaria del Amazonas, Trapecio Amazonico, Rio Lore- 
toyacu. Alt. about 100 m. “‘Bush. Flowers white.’’ October, 1945. 
R.E. Schultes 6727.—November 2, 1946. G.A. Black et RE. Schultes 
46-295. 

The Tikuna Indians and the Brazilian rubber tappers 
of the Trapecio Amazénico assert that the seeds of this 
bush or small tree are extremely dangerous when eaten. 


[ 154 ] 


Prater XLIL 


® saccifera (Mart) 


Nz 


id 
2 


, fruit dissected, approximately 5 


3 


Land 2, habit, approximately 3 >. 


ged. 


greatly enlar 


5S 


4, section of leaf 


Duroia saccifera (Mart.) Hooker fil. ex K. Schu- 
mann in Martius Fl. Bras. 6, pt.6 (1889) 862, t.146, fig. 1. 

CotombBra: Comisaria del Amazonas, Rio Apaporis, Soratama and 
vicinity. ‘‘Tree 10 m, tall.’’ March 26, 1952. R.E. Schultes et I. 
Cabrera 16065, 


The natives in the middle course of the Rio Apaporis 
state that the seeds of this small forest tree are toxic 
when ingested. 

This collection extends the known range of Duroia 
saccifera into the Amazonian basin of Colombia; it has 
hitherto been known from Amazonian Brazil and the 
Rio Guainia area of Venezuela. 


Duroia Sprucei Rusby Descr. New Sp. S. Am. PI. 
(1920) 133. 

CoLombia : Comisaria del Amazonas, Trapecio Amazoénico, Rio Lore- 
toyacu, “Small tree. Flood bank.’’ May, 1946. R.E. Schultes 7127. 


—Rio Apaporis, Soratama, above mouth of Rio Kananari. June 18, 
1951. RE, Schultes et I. Cabrera 126465. 


The seeds of this small tree have the reputation 
amongst the natives of the Rio Apaporis of being poison- 
ous to eat. 

The type of Durota Sprucei was collected in Vene- 
zuela. ‘The collections cited above extend the known 
range of the species significantly southwestward. 


Psychotria L. 

So far as the available chemical literature is concerned, 
there is little if any indication that the seeds and fruits 
of species of Psychotria would be toxic when ingested 
by man. Reports to this effect, however, are so frequent 
and are scattered over such a wide area that phytochemi- 
cal studies based upon these ethnobotanical observations 
are justified. 


[ 156 | 


PLATE 


? 


G7 
~ ys aes 
: rE. ¥2 
baa cao Ms aie 

4 Oia 
4, 


Fletcher 


oN 


—==pe A 


DUROIA 


, ¥ petiolaris 


% P 
se Hook. fil. 
ql 
h. 
Mi " \ - 
2, flower in bud, approximately 34 *. 3, flower 


Aa 3 
SK 


> ~ 
oe 5 om 


habit, approximately 
opened, approximately 36 


Psychotria carthaginensis Jacquin Enum. PI. 
Carib. (1760) 16. 

CoLtompia;: Comisaria del Amazonas, Rio Apaporis, Raudal Yaya- 
copi (La Playa) and vicinity. Quartzite base. Alt. about 800 ft. 
‘Bush. Flowers white.’’ April 15, 1952. R.E. Schultes et I. Cabrera 
16203,—**Bush. Fruit red. On rocks of bank. Makuna name wy- 
soo-dé.”’ August 18, 1952. Schultes et Cabrera 16890. 


The Makuna Indians assert that the fruits of this 
shrub, if eaten, cause a poisoning that results in several 
days of weakness, fevers, nausea and disturbed vision. 


Psychotria involucrata Swartz Prodr. Veg. Ind. 
Occ. (1788) 45. 

CotomsBia: Comisaria del Amazonas, Rio Amazonas, Leticia and 
vicinity. September 1946. R.E. Schultes 8231.—Rio Apaporis, Sora- 
tama, between Rio Pacoa and Rio Kananari. “‘Bush, Highland.’’ 
August 20, 1946. R.E. Schultes et I. Cabrera 18629. 


The Tikuna Indians of the Leticia region and the 
Makunas of the Rio Apaporis both consider the fruits 
of Psychotria involucrata poisonous when ingested. 


Psychotria nudiceps Standley in Field Mus. Publ. 
Bot. 8 (1981) 378. 

CoLtompia: Comisaria del Amazonas, Trapecio Amazénico, Rio Lo- 
retoyacu. October 1946, R.E, Schultes 8408. 


[t is acommonly accepted belief amongst the rubber 
tappers in this part of Colombia and adjacent Peru and 
Brazil that the fruits of Psychotria nudiceps are toxic 
when eaten. 


Psychotria psychotriaefolia (Seem.) Standley in 
Contrib. U.S. Nat. Herb. 18 (1916) 133. 


Cotompra: Comisaria del Putumayo, Mocoa and vicinity. “‘Leaves 
used with yajé.”’ No date. C. Naranjo s.n. 

Ecuapor: Provincia del Napo, Rio Aguarico, Dureno. Alt. about 
800 ft. ‘‘Area near river and flood region. Shrub leaves and fruit 
used in preparation of the yajé narcotic as an admixture with Banis- 
teriopsis.’’ Alt. 800 ft. May 29, 1966. H.V. Pinkley 225; 235. 


[ 158 ] 


PuaTE XLV 


i ~ 


WV 


LOE LP 
Ale 

“fe PSYCHOTRIA 
nudiceps 


Stand]. 


ih 


S4 of /. ae - 
Wie es, 
Fy ce io 


SSS 


iy << 
oS Be 


PSYCHOTRIA 


carthagenensis 


Ny EAA pk 
iy ay {27 KS 


Jacq. Vey 


1 and 2, habit, approximately 4X. 


[ 159 | 


PiatE XLVI 


yt yy Uf - 
‘| V, y 


Wy py) Dh / J Witorg 
ims within sibiond 
I ( 


if i 
SER 


- 
ee 


Wh id 


Ip 


We, 


wie 


Flek her 


PSYCHOTRIA 


involucrata Sw. 


INs 


1 and 2, habit, approximately $ . 3, dried inflorescence, approximately 35 >. 


[ 161 ] 


- 7 


PuaTE XLVII 


PSYCHOTRIA foo y 


psychotriaefolia | . ge 
(Seem.) Standl. | Bee ¥ 8 


Ba of 


Habit, approximately natural size. Flower, approximately 5 *. Dissected 
corolla, approximately 5 . Fruit, approximately 4 *. Cross section of fruit, 


approximately 4} . 


163 ] 


The earliest report of the use of a Psychotria as an 
additive to ‘‘fortify’’ the hallucinogenic yajé drink pre- 
pared basically from Banisteriopsis Caapi or B. inebrians 
was made in 1967 (Schultes, R. E. in Efron (ed. ) ‘‘Ethno- 
pharmacologic search for psychoactive drugs’’ (1967) 51). 
At that time, the identification could be made only to 
genus. Now, asa result of the field work of Mr. Pinkley 
amongst the Kof’n Indians, it is possible to assign the 
species with certainty to Psychotria psychotriaefolia. 
Pinkley has, furthermore, been able to determine the 
sterile collection Naranjo s.n. to this species. 

In the morning, after one of the all-night yajé-sessions 
which he attended amongst the Kofins, Pinkley found, 
in the dregs at the bottom of the clay pot which had 
held the narcotic drink, a sediment containing leaves of 
Banisteriopsis and small rubiaceous fruits. These dregs 
might not have attracted the notice of an untrained ob- 
server. Since, however, Pinkley was especially interested 
in the complexities of yajé-preparation, these specimens 
raised his curiosity and, pursuing the problem ethnobo- 
tanically, he established the fact that Psychotria psycho- 
triaefolia is not an uncommonly used additive in this 
region. 

The Kofins call Psychotria psychotriaefolia by the 
name o-pri-to. One of Pinkley’s interesting observations 
indicates that these Indians refer to the ‘‘heavenly 
people’’, with whom they commune during the yajé in- 
toxication, by the same term o-pri-to. He further re- 
ports that Mr. M.B. Borman, a missionary-linguist 
studying the Kofiin language, states in a letter that both 
Banisteriopsis Rusbyana and Psychotria psychotriaefolia 
are added to the yajé drink prepared from B. Caapi to 
“increase the visions and make them of longer duration’’. 

‘These observations serve to emphasize long felt urgen- 
cy for a thorough chemical study of Psychotria and very 
closely related genera, such as Palicourea. 

The collections Pinkley 225 and 235 were identified 
by Dr. John D. Dwyer of the Missouri Botanical Garden. 

[ 164 ] 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


Vor, 22, NaS 


CampripGe, Massacuuserts, June 13, 1969 


TREE DATURA DRUGS OF THE 
COLOMBIAN SIBUNDOY' 
BY 
ME.LvIN L. BristTo.’ 


IN southernmost Colombia high on the eastern flank of 
the Andean cordillera lies a small and isolated montane 
basin, the Valley of Sibundoy (Plate NLVIII). Several 
thousand Kamsi-speaking Sibundoy and three Inga- 
speaking groups have inhabited the Valley for several 
centuries, perhaps for much longer.” The collapsing agri- 
cultural terraces clearly visible at many places on the 
valley sides indicate a populous pre-hispanic occupation, 
and the Sibundoy believe that their ancestors have lived 
in the Valley from very early times. As yet, however, 
there is no evidence linking the early terrace builders 
with the Sibundoy or with any other native group in 
southern Colombia. 

Despite the apparent isolation of the Valley of Sibun- 
doy, the natives have probably always been in contact 
with a diversity of other aboriginals. Today, as in the 
Sixteenth Century, there are three trails leading out of 
the Valley to the east, west and north. Two of these 
have recently been supplanted, for a road now connects 
the highland capital of Pasto twenty miles to the west 


‘Received for publication January, 1968. 
2H. L. Lyon Arboretum, University of Hawaii, Honolulu, Hawaii. 
* Inga is one of the northernmost Quechua dialects (9, 24). 


[ 165 ] 


with Mocoa in the eastern lowlands. The third, and 
shortest trail, connecting the Valley with the Rio Juan- 
ambu drainage to the north, is probably little changed 
from 1801 when Humboldt and Bonpland traversed it 
twice, or from 1541 when Hernan Pérez de Quesada 
fled homeward along it from his wretched search for 
El Dorado. 

During the long period of relative isolation, a great 
variety of curious cultivated plants were brought into 
the Valley. Some are of scant importance today and may 
never have enjoyed a wide appreciation among the Val- 
ley’s inhabitants. Others, the predominant food, medi- 
cinal and narcotic plants, have come to assume very great 
importance in the economic and social life of the natives. 
Certain plants, known nowhere else, have evolved in the 
Valley under the influences of cultivation. Such has 
come to pass with the tree Datura drugs. 

The genus Datura consists of eight to twelve herba- 
ceous species (2, 8, 28), with their centre of diversity 
in Mexico and southwestern United States: and three 
or more (7) to fourteen (28) arborescent species centered 
in the northern Andes. In the absence of a modern and 
comprehensive revision of the genus, Safford’s account 
of 1921 has been widely accepted. However, were we to 
continue employing his species concept today, the diver- 
sity of herbarium material available now would allow us 
to define thirty or more ‘‘species’’ of tree Daturas (sect. 
Brugmansia) alone. Recently, I have pointed out that 
almost all tree Daturas belong to one of three species 
and that the few remaining plants are probably hybrid 
or aberrant individuals (7). The variability expressed in 
the tree Daturas as a group has been enhanced through 
their cultivation by many native peoples; in fact, their 
absence from any natural vegetation implies that their 
recent evolution has taken place entirely under man’s 
influences. 

| 166 | 


PuateE XLVIII 


COLOMBIA Valley of Sibundoy 


Comisaria Especial del Putumayo, Colombia 
ROAD ——— PASTURE 
TRAIL ------ MARSH 


! 


\ Sibundoy 


‘ at: CAD 
 SIBUNDOY = -§ . 
,\ OU —— 
as 5 
r § [ TRIBAL. - oe 
. ~ ae 4 re To Mococ 
“a, MN / es 
% (G y ~R. Putumayo 
ee 5 AREA J ~ San _ 
¥ ad A Francisco 
if ‘ zu L >> 
- ae MEG 
= , ee 
\ > ie 
| sa 
\ 
Al wlues Pe 
\ Ps 
a ae 
y eeet <_ se 21a8% Juan Yunguilloe 
: . ‘ 
N - 4 ~~ 
r 
Sibundoy 
” ~ £ a . ae 
x a ¢ ° 
}! rn ; = 4 "e :] R 
“y oe 
g ¢ IC 20 Kms 
Trall : l | 
Road 2 
fe) ' 2 3 4 Kms 76°55 
a L 1 Lo 


Valley of Sibundoy, showing the area inhabited today by the Sibun- 
doy Indians. (Areas at the western and southwestern margins of the 
Valley inhabited by Colén, Santiaguefio and Sanandresefio Indians 
are omitted. ) 


In the Valley of Sibundoy, most of the Daturas be- 
long to the species Datura candida (Pers.) Saff., though 
D. sanguinea R. & P. is also present. They are much 
planted by the natives for ornament in hedge rows, in 
house yards and in the multi-purpose gardens; and the 
abundance of huge, white flowers which appear four 
times annually make them the principal visual focal point 
everywhere. Containing atropine and hyoscyamine, these 
borracheras (‘inebriants’) have been used variously by the 
natives both as psychotropic and medicinal preparations. 
While one of the cultivars, ‘Buyés’, is frequently seen 
throughout the inhabited parts of the Valley, most of 
the cultivars are encountered only very infrequently. So 
far as [am aware, the unique Daturas of the Valley were 
not known to outsiders until Hernando Garcia- Barriga 
found them in 1985. 

Frequent suggestions that the Sibundoy tree Daturas 
are infected by viruses has focused attention on this as a 
major cause of the peculiar leaves by which several of the 
cultivars are recognized (8, 22, 85, 36). However, early 
in the course of my thirteen months of observations in the 
Valley in 1962-68, I realized that each cultivar is geneti- 
cally distinet, quite apart from the possible influence of 
viruses on the leaves. Differences among the cultivars 
which I believe to be genetic lie in the morphology of the 
flowers and fruit, in the incidence of chromosomal inver- 
sions and in the amount of aborted pollen. Some of the 
unique leaves are also reflections of genetic uniqueness 
and not of virus infection. In recognition of ethnomedi- 
cal, pharmacologic, chemical and phytopathologic inter- 
est in the Sibundoy tree Daturas, an important objective 
of my study has been to understand their morphologic 
and cytologic variability, to distinguish among them, 
and to define them. 


[ 168 ] 


Variation in Datura candida ‘ Buyés’ 

Most specimens of Daturas in the Valley of Sibundoy 
are called buyés borrachera by the Kamsa-speaking Sibun- 
doy, or D. candida ev. Buyés as described below. They 
represent several, or possibly many, genetic lines, as 
evidenced by marked differences in the size and shape of 
the flowers. At the present time, however, the natives 
are little, if at all, aware of these variations, and they do 
not discriminate verbally within their concept of buyés 
borrachera. For later purposes of comparison, it will be 
helpful here to record the limits of variation of D. can- 
dida ‘Buyés’ in the Valley of Sibundoy. To this end, a 
representative sample of leaves and flowers was taken 
from twenty-eight trees scattered throughout the area of 
the Valley inhabited by the Sibundoy. With the addition 
of other collections, there are thirty-seven trees repre- 
sented, to give an indication of the range of variability of 
this cultivar. Table I gives the range of variation in D. 
candida ‘Buyés’ for six of the eight characters found to 
be most useful in distinguishing the nine Sibundoy culti- 
vars of D. candida. 


7 7 A . ‘ ~.9 ‘i 
Taste I. Range of variation in D. candida Buyés’ (meas. in mm.). 


median 
Leaf ratio (W/L) .346-.577 .460 
Calyx length 85- 161 128 
Corolla length 196— 293 253 
Stamen length (including the 132- 199 165 

adnation of filament to corolla) 

Anther length 30- 41 35 
Pistil length 143- 212 Lie 


The most conspicuous feature of these trees is certainly 
the brilliant white corolla, but despite this, its variation 
in size has no formal recognition among the Sibundoy. 
In many flowers, the edge of the corolla between the 
lobes is emarginate, but in some it is straight or well 


[ 169 ] 


rounded. Counts of aborted pollen in six plants ranged 
from 20.7 to 80.4 per cent. 

D. candida ‘Buyés’ is searcely distinguishable from 
many examples of D. candida in widely separated areas 
of tropical America. In the characters studied, its range 
of variation is less than that of the species as a whole. 
The rare Sibundoy cultivars of this species have never 
been found outside of the Valley of Sibundoy, and it is 
my assumption that they are all derived, at least in part, 
from D. candida ‘Buyés’.* 


Chromosome nu mbers 


Heiser (16) found the chromosome number of JD. can- 
dida as represented by the collection Heiser 6119 (IND), 
to be n=12, and Barclay (2,3) found 2n=24 in D. can- 
dida ‘Culebra’ (Barclay & Schultes 286) and in D. vul- 
canicola A.S. Barclay. 

Preparation of aceto-carmine stained microspore 
mother cells permitted observation of the chromosomes 
after 48-72 hours. The chromosome counts for each cul- 
tivar were made on either one or two plants as indicated 
by the voucher collection numbers in Table II. Seven of 
the nine D. candida cultivars were examined, and all 
were seen to havea chromosome complement of n=12. 
All counts were made after metaphase I, most at or 
following metaphase II. The data are summarized in 
Table IL. Figure la, illustrates the chromosomes of 
D. candida ‘Culebra’. 

Twelve is the haploid chromosome number of Datura 
sanguinea R.& P. (7, 16), of *‘D. suaveolens’’ (15), and 
of all the herbaceous species of Datura (1). 

‘TI recently encountered a tree referable to D. candida ‘Amar6n’ 
near Las Cruces in southernmost Narifo. Thus the question is raised 
as to whether this was brought from the Valley of Sibundoy, or 


whether it is part of an early population of plants from which one or 
more of the Sibundoy cultivars might have originated. 


[ 170 | 


PLaTE XLIX 


a 4 é j i . : : ¢ " 3 


Datura candida (Pers. ) Saff. ev. Buyés. (Upper) A massive display 


of flowers. (Lower) Habit of the tree, growing in a cornfield. 


Taste Il. Chromosome numbers of D. candida cultivars, 


No. of cells during microsporogenesis with Voucher 

Cultivar n=10 n=11 n=12 n=18 collection* 
*Buyés’ l Q 137 3 1117, 1266 
‘Ocre’ - 5 57 - 1267 
*Biangan’® l 3 43 - 1431 
*Amaron’ - _ 29 1 564 
‘Quinde’ l l 30 - 1304, 1483 
*Munchira’ 1 8 46 l 1268 
‘Culebra’ - 6 122 5 1112, 1400 


* On deposit at the Economic Herbarium of Oakes Ames, Harvard Univer- 
sity, and the United States National Herbarium. 


Anaphase bridges 


In four cultivars of D. candida, the frequency of ana- 
phase [ bridges in pollen mother cells was recorded. ‘The 
formation of the anaphase bridge leads to uneven distri- 


0 
sy 
QO 

? 


a b 


Figure 1. Chromosomes of D. candida ‘Culebra’. (a) Metaphase I 
(=< 500). (b) Anaphase II bridges ( * 2000). 


bution of chromosomal material between the first division 
meiotic products and to consequent imbalance in the four 
resulting microspores. ‘Table I1] summarizes the data 
and includes frequencies of pollen abortion from ‘Table 
LV for comparison. 

In ‘Ocre’, no pollen mother cells were seen with more 
than one anaphase I bridge. *‘Buyés” plants showed 
PMC’s with one to four bridges, while in ‘Culebra’ 
there were up to nine bridges between first division 
nuclei. Figure 1b, illustrates an occurrence of anaphase 


Tasre III. Anaphase I bridges formed during microsporogenesis 
in D. candida cultivars. 


Total No. PMC’'s % PMC’'s observed % 

PMC’s with A I with A I abnormal 
Cultivar observed bridges bridges pollen 
*Buyés’ 213 35 16.4 ‘e.A* 
‘Ocre’ 358 64+ 17.9 50.8 
*Amaron’ 1000 30 3.0 32.8 
‘Culebra’ 537 97 16.9 83.3 


**Buyés’ trees eand fonly. 


bridges in ‘Culebra’. The chromosomal irregularities 
leading to the formation of anaphase bridges are un- 
known. In many cases the bridges were accompanied 
by one or more fragments, implying that chromosomal 
inversions are responsible for some, but not all of the 
observed bridges. 


Pollen abortion 


Preliminary examination of the pollen of D. candida 
‘Culebra’ disclosed a predominance of small, either 
spherical or misshapen, spores among the large and well 
sculptured ones. I decided to investigate for compara- 
tive purposes the frequencies of such abnormal pollen in 
all of the D. candida cultivars. In all but two cases, a 
minimum of two widely separated trees was examined, 
one by myself and one by an assistant. The results of 
these examinations are recorded in Table IV. 

It can be seen that the production of abnormal pollen 
ranges from a low of 5.7 percent in ‘Dientes’ to a high 
of 83.8 per cent in ‘Culebra’. As had been thought from 
purely morphological evidence, the cultivars ‘Buyés’ and 
‘Dientes’ do not consist of a single clone each; with both 
groups there are widely differing capacities to produce 
normal pollen. 

Some of the abnormal pollen may be accounted for by 
the inequality of meiotic products resulting from ana- 


[ 173 | 


PLatTE L 


Anthers | | 
Pistil 
Stamens | 
Corolla 
Calyx 


| | | 


‘Amarén' 'Salaman' ‘Quinde’ 


[ fo of | 
ia ae 


‘Munchira’ ‘Culebra’ ‘Andrés’ 


Ranges of variation in the lengths of five floral parts of Datura cul- 
tivars compared with the respective ranges of variation in D. candida 
*Buyés’. The values for ‘Buyés’, from Table I, are standardized to a 
common size in the uppermost ideogram, and values for the other cul- 
tivars (cf. taxonomic treatment) are expressed in proportion to these. 


Taste 1V. Frequencies of abnormal pollen in D. candida cultivars. 


Total % 
pollen abnormal abnormal 

Cultivar Tree — observed pollen pollen 
‘Buyés’ a 500 142 

b 270 770 56 198 25.7 

C 500 208 

d 500 1000 216 424 42,4 

e 700 535 

f 500-1200 402 937 78.1 
‘Dientes’ a 700 39 

b 529 29 

c 500 1729 30 98 bit 

d 500 129 25.8 

e 518 176 34.0 
No. 7111 a 600 260 43.3 
‘Ocre’ a 608 308 

b 500 1108 255 563 50.8 
*Biangan’ a 800 472 

b 564 370 

c 500 1864 299 1141 61.2 
*Amaron’ a 764 O25 

b 700 1464 255 480 32.8 
“Salaman’ a 1546 173 11.2 
*Quinde’ a 594 175 

b 500 211 

c 500 199 

d 500 2094 180 765 36.5 
*“Munchira’ a 500 29] 58.2 
‘Culebra’ a 615 506 

b 490 416 


c 250 1355 207 1128 83.3 


phase bridges as noted above in Table III. ‘The remain- 
der might well be caused by the presence of genetic fac- 
tors for pollen abortion; Blakeslee and his co-workers 
found thirty such factors in their extensive cultures of 
herbaceous Daturas (1). 


Flower variation among the Datura candida cultivars 


All of the cultivars have markedly similar flowers, with 
the exception of the bizarre ‘Culebra’ which is discussed 
later. Nevertheless, each displays a distinctive range of 
variation in the length of the several floral parts. Plate 
L. compares nine of the cultivars according to the lengths 
of five of their floral parts. It is evident that with but 
few exceptions the ranges of variation of each cultivar 
are either wholly within, or largely overlapping, those 
of D. candida ‘Buyés’. This strongly suggests that most 
of the cultivars were selected from the large and variable 
population of D. candida ‘Buyés’. It further suggests 
that the Sibundoy did not select and propagate new 
forms (cultivars) characterized by floral differences, but, 
as we shall see below, they selected leaf differences for 
preferential maintenance as clones. 


Fruit variation 


Most of the fruits found in prolonged searching 
throughout the Valley were on ‘Buyés’ and ‘Amaron’ 
trees which are by far the most numerous. Some fruits 
were also seen and collected on ‘Dientes’, ‘Quinde’ and 
‘Biangiin’ trees; the remainder of the D. candida culti- 
vars are believed by the natives never to form fruit. The 
indehiscent fruits eventually dry up, and the pericarp 
disintegrates slowly: but the seeds are not released, and 
the dried fruit often remains on the tree for several 
months. Plate LI shows the variation in fruit size and 
shape among the five cultivars known (however rarely) 


[176 ] 


Piate LI 


‘doryeur[dxa Jo} 4x93 9as * UOIYVMTY —VW ‘(,s9dng) 
DIJ{[ pue (,sequsiqd,) L777 ‘(,apurng), ) T@rE *( upsueig |) COLT iSuoryat[oo Sulmoyypoy 
94} Wolf aiv ‘azis [Rinjeu F uaMOoYs *s}inly 94], “SIBALQ[ND Dpipuyo “CT UL UOTZBLIVA YINI 


YU} PIM 


ee 
OL 
- ; ’ 7 x. a ’ J —_— 
ae ee eee ee 
, oe ae oe ee ee 
sghng. sajuald, apuind, upbueig, | 3 


to produce fruit. Most of the nine ‘Amar6n’ fruits col- 
lected fall within the ‘Buyés’ variation pattern; two are 
the same as ‘Quinde’. A single ‘Buyés’ tree (Bristol 
1109, 1193) has unusually long (11.7-24.7 em.), more 
narrow fruits which are not included in Plate LI. Meas- 
urements were made on ripe fleshy fruits. 


Leaf variation 


Despite the highly conspicuous and attractive flowers 
of all the cultivars, the leaves are the centre of interest 
to the natives, to whom they are the structures of prin- 
cipal economic importance. Plate LII depicts a repre- 
sentative leaf of each cultivar. The striking malforma- 
tion in four of the cultivars has led to statements that 
the plants are diseased, probably by viruses, or that they 
are ‘‘ ‘virus-races’ *” (8, 22, 85, 36). 

Recently, Robert Kahn and R. Bartels have isolated 
a previously unknown virus from several of the cultivars, 
including ‘Buyés’ which never has erose or otherwise 
malformed leaves (21). These investigators have named 
the virus Colombia Datura Virus (CDV) and report that 
its symptoms in Datura are leaf distortion, mottling or 
yellowing, but that sometimes there are no symptoms. 
Leaves of the common D. candida ‘Buyés’ in the Valley 
of Sibundoy are never distorted, and mottling and yel- 
lowing, if present, are rare. 

It seems probable that there is no direct causative re- 
lationship between CDV and leaf shape in the D. can- 
dida cultivars. At the same time, the genetic control of 
leaf shape is enigmatic, for careful examination of the 
irregularly leaved cultivars will disclose on every plant 
a few small to medium sized ovate and entire leaves, as 
in “Buyés’. These leaves invariably occur on one or two 
branches which are growing more rapidly than all others 
on the plant. Given this exception, it is tempting to pos- 


[ 178 ] 


PLATE LII 


Representative leaves of all Sibundoy Datura cultivars, with collec- 
tion numbers indicated: 1, ‘Amar6n, 564; 2, ‘Guamuco’, 1420; 3, 
‘Salaman’, 1432; 4, ‘Sangre’, 1309; 5, ‘Quinde’, 1333; 6, ‘Andrés’, 
1314; 7, ‘Culebra’, 1112; 8, ‘Munchira’, 1268; 9, ‘Buyés’, 1388; 
10, ‘Biangdn’, 890; 11, ‘Dientes’, 1447; 12, ‘Ocre’, 1267. 


tulate that the highly irregular formation of the leaves 
is caused by a critical concentration of a diffusable sub- 
stance, and that in the exceptional cases of very vigor- 
ously growing stem apices the critical level of concentra- 
tion is not reached. Such a substance might be viral or 
hormonal in nature. That the natives find the smallest, 
most malformed leaves the most toxic suggests that even 
the tropane alkaloids should not be ruled out of con- 
sideration as possible inhibitors of regular leaf develop- 
ment. As Robert Kahn has suggested to me, grafting 
experiments should help to clarify the development of 
the various leaf shapes. 


ae 
Seedlings 


Through thirteen months of constant observation of 
the Datura cultivars in the Valley of Sibundoy, no seed- 
ling tree was seen. It has been possible, however, to 
grow healthy seedlings of three clones, ‘Buyés’, ‘Ama- 
ron’ and ‘Quinde’, in the greenhouses of the Biological 
Laboratories of Harvard University. The seedlings 
within each cultivar are not uniform. Seeds collected 
trom D. candida ‘Quinde’ trees have produced seedlings 
the leaves of which resemble ‘Dientes’ (1596, 1611; 
ECON) as well as ‘Quinde’ (7607; ECON). Seedling 
variation from a single ‘Quinde’ fruit includes both 
‘Quinde’ and ‘Dientes’ types (7608; ECON), suggesting 
that a ‘Dientes’ tree may have been the pollen parent. 

‘Quinde’ seedlings exhibit the same failure to form 
‘Quinde’-type leaves on a very vigorously growing stem 
that was observed on mature trees in the Valley of Sibun- 
doy. As the growth rate of the seedlings slows some- 
what, these same stems that grew vigorously produce 
typical ‘Quinde’-type leaves. Assays of these seed grown 
‘Quinde’-type leaves for the presence of mechanically 
transmittable virus by Robert Kahn (pers. comm.) have 


[ 180 | 


been negative, as would ordinarily be expected. ‘Thus 
the formation of ‘Quinde’-type leaves by apparently 
healthy seedlings implies that the typical leaves of 
‘Quinde’ trees are formed independently of the virus 
(CDV) infecting them. 

One may compare the successive occurrence of differ- 
ently shaped leaves to the widely encountered phenome- 
non of juvenility, but in so doing it must be noted that 
only four of the D. candida cultivars are involved, while 
the other five are not. I have not seen any descriptions 
of juvenile leaves in the genus Datura. 

Blakeslee found a variant of D. Stramonium with a 
leaf similar but not identical to that of ‘Quinde’ (1). He 
named these ‘Quercina’ and soon established that their 
unusual leaf shape depended upon asingle recessive gene. 
Curiously, he also discovered that the juices of the plant 
would transmit the characteristic to normal JD. Stramo- 
nium, and he believed that the Quercina effect was caused 
both by a gene and by avirus. Further studies of the sup- 
posed virus have not been undertaken to my knowledge. 


Geographical centre of dwersity 


More variation in Datura candida is found in the Val- 
ley of Sibundoy than is known at any other locality. 
Such acentre of diversity in an isolated valley apparently 
inhabited by ancient agriculturists suggests the possibili- 
ty of domestication and early evolution here. However, 
that the Valley of Sibundoy is not the centre of origin 
of any tree Datura is attested by the absence of all Da- 
tura spp., and any solanaceous plants resembling them, 
in the non-cultivated flora. Were the Valley the site of 
domestication, we would expect to find some plants capa- 
ble of self-reproduction, either sexually or asexually, and 
not wholly dependent upon man for survival. Evidently 
one or more clones of D. candida were brought into the 


[181 ] 


Valley at some remote time. Today, J. candida is found 
neither in the eastern lowlands, nor in the western high- 
lands around Pasto, but several plants of D. candida 
‘Buyés’ (Bristol 1785) were seen in the upper Rio Juan- 
ambu drainage to the north. It may be postulated that 
D. candida was introduced to the Valley of Sibundoy 
via the trail from the Juanambu, and that following one 
or more such introductions the Valley became a centre 
of diversity. 


b em , 
Propagation 


All of the Datura cultivars in the Valley of Sibundoy 
are propagated exclusively by man through large cut- 
tings. During the annual clearing of the gardens in 
preparation for the planting of maize and beans, some 
of the many trees and shrubs in the gardens are pruned, 
pollarded or even cut back to the ground. The removal 
of branches from a Datura plant to permit more planting 
space is the usual occasion for propagation. A branch 
measuring half a meter or more is simply thrust into the 
loose sandy soil and given no further attention. Occa- 
sionally, a more methodical planting is undertaken when 
the branches are cut into fairly uniform pieces and 
planted out in a row as a kind of hedge along the edge 
of a garden. These plants will be weeded during the 
general garden weedings four times a year. 

Tree Datura branches at different stages of maturity 
differ in their capacity to root easily from cuttings. All 
of the leaves are borne on herbaceous stems which are 
only tardily lignified. Occasionally, usually after heavy 
pruning or pollarding, one or more stems grow up with 
unusual vigor and often reach a diameter twice that of a 
normal stem. The lignified part of normal stems is the 
least capable of forming roots under casual treatment; 
but the occasional, thick and uncommonly vigorous stems 


{ 182 | 


L 4 


develop roots quickly and abundantly when cuttings are 
taken from the region which has begun to lignify. One 
clone, *Munchira’, is difficult to propagate, since its stems 
are typically undersized, and the few stunted plants in 
existence are only rarely cut back. When this is done, 
however, the expected growth of thick, vigorous shoots 
appears, and cuttings made from these root easily. 


Abundance of Datura cultivars 


In the Valley of Sibundoy, tree Daturas are more con- 
spicuous and more abundant than at many other localities 
in southern Colombia. At least 60 per cent of the trees 
are D. candida *Buyés’, while another 80 per cent are 
D. candida *Dientes’ and *Amaron’. All of the eeonomi- 
‘ally more important cultivars comprise the remaining 
10 per cent. ‘Table V indicates the estimated number of 
individuals of each cultivar in 1968, and emphasizes the 
rarity of many. 

Taste V. Estimated number of individuals of Datura cultivars 

in the Valley of Sibundoy in 1963. 


ca. 1000 *Buyés’ 

ca, 300 *Dientes’ 

ea, 200 *Amaron’ 
‘Culebra’ 
*‘Quinde’ 


less than 40 re ; 
Guamuco’ 


‘ . , . 
Biangan 


‘ e . 
Munchira 
less than 20 ba 
Sangre’ 


‘Andrés’ 
less than 10 ‘Ocre’ 


to , 
Salaman’ 


It is difficult to judge whether the relative scarcity of 
nine of the cultivars reflects their more recent appearance 
in cultivation, or the degree of jealousy with which they 


[ 183 ] 


are guarded from other potential owners. A medicine- 
man once stated that he could not afford to be seen near 
an unusual Datura in another man’s garden (6). [If such 
a prohibition is general among the Sibundoy, the tree 
Daturas, however useful, may be increasing at an ex- 
tremely slow rate. The two cultivars thought by the 
Sibundoy to be the most toxic, J. candida *Munchira’ 
and ‘Salaman’, are among the least abundant. 


Common names 

The three linguistic groups of people in the Valley of 
Sibundoy agree in grouping all the Datura cultivars in 
the Spanish term borrachera (‘inebriant’).” However, 
the Sibundoy believe this to be a Kamsa morpheme, not 
a loan word. Ten of the twelve cultivars are discrimi- 
nated verbally, but because of the rarity of several, very 
few Sibundoy can name all ten. To name all ten, one 
must draw words from three languages, but because of 
generalized borrowing among these three, a speaker may 
not be aware of this. Kamsa lacks names for three cul- 
tivars, Inga for two. Table VI gives the common names 
recorded in 1962-63. A full account of the vernacular 
names is given with the taxonomic treatment of the cul- 
tivars. My capabilities did not permit the most precise 
form of phonetic records. Accurate pronunciation will 
be approximated most frequently within the Spanish, 
not the English, phonetic framework. An exception is 
bui-ish which | have written here buyés, conforming with 
the notation of Juajibioy (19). 


gconomue importance 


Both my own investigations and the reports of others 
indicate that the Daturas are used about equally for 


’ Several non-solanaceous, non-psychotropic plants are also called 
borrachera, e.g., lresine celosia L., I. herbstii Hook.f., and other un- 
identified plants. 


[ 184 ] 


Tas_te VI. Common names employed in the Valley of Sibundoy 
for Datura cultivars. 


Common name Language Scientific name 
1. Amarén* Spanish D. candida “Amaron’ 
2. Andaqui Inga Datura ‘Andrés’ 
3. Biangdén* Inga D. candida *Biangan’ 
4. Borracherat Kamsa D. candida; 


(Spanish) D.candida ‘Buyés’, “Dientes’ 


5. Borrachera de agua Spanish  D. candida ‘“Buyés’, ‘Dientes’ 

6. Buyés* Kamsia D. candida ‘Buyés’, “Dientes’ 

T. Cae Inga D. candida ‘Quinde’ 

8. Cucu Inga D. candida ‘Amaron’ 

9. Culebra* Spanish D. candida ‘Culebra’ 

10. Chontaruco Inga D. candida * Biangan’ 

11. Danta Inga D. candida ‘Biangan’ 

12. Floripondio Spanish  D.candida “Buyés’, ‘Dientes’ 
13. Guamuco* D. sanguinea ‘Guamuco’, ‘Sangre’ 
14, Guamuco blanco Spanish  D.candida ‘Buyés’, ‘Dientes’ 
15. Guamuco floripundo D. sanguinea ‘Guamuco’, ‘Sangre’ 
16. Ngunsiana Kamsa D. candida ‘Quinde’ 

17. Lengua de tigre Spanish  D.candida ‘Culebra’ 

18. Mutscuai Kamsa D. candida ‘Culebra’ 

19. Munchira* Inga D. candida “Munchira’ 

20. Quinde* Inga D. candida ‘Quinde’ 

21. Salamén* P D. candida ‘Salaman’ 
22. Salamanga P D. candida ‘Salamin’ 

23. Salvanje Kamsa D. candida ‘Salaman’ 


* Most frequently employed while speaking Spanish. 
t Borrachera is usually ‘tree Datura’. When the context is not clear 
it follows the cultivar name,e.g.,culebra borrachera snake tree Datura’. 


medicinal and for psychotropic preparations by the 
Sibundoy. This information is summarized in ‘Table 
VII. Several of the cultivars are relatively unimportant, 
but to four attaches an aura of great respect for their 
power to alter consciousness drastically — even to cause 
death. 

Most of the Sibundoy indicate a genuine interest in 
the borracheras, but individuals vary greatly in the ex- 


[ 185 ] 


- Table 7. 


Uses of Datura cultivars. 


(x indicates one reports, ) 


Internal 
Cultivar a 
2. 
°o 
fs 
» 
° i 
a (o) 
oO ” 
e | 3 
oy Ou 
1. Guamuco } 
2. Sangre 
3. Buyés 
Le orenvea| 
S. Ocre 
6. Biang4n x x 
xl 
x2 
7. Amarén 
8, Salamin x 
3. Quinde xxx 
xxx 
10, Munchira xxx 
- x 
x 3 
xh 
ll. Culebra 
x 5,6 x5 
x7 
x 8 x 8 
x9 
x 10 
x il 
12, Andrés | x 


Use 
External 
Medicine 
oO 
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* “*Report’’ here means one of the following: (a) one Sibundoy’s personal 


footnoted. 


1. Idrobo 2231 (COL) 


2. (38) 

Idrobo 2223 (COL) 
(20) 

Schultes 3256 (COL, ECON,GH) 11. 
(29) 


Schultes 20079 (GH) 


7. 

8. (35) 

9, Idrobo 222 
10, 


1 (COL) 
Fernandes 2641 (COL) 
Mora 1023 (COL) 


report to me, (6) one herbarium specimen notation by another botanist, (c) one 
primary literature statement not duplicating a herbarium notation. 6 and care 


tent of their knowledge. For example, very few, perhaps 
only the medicine-men, are able to name all the cultivars 
accorded recognition by the culture; but even some of 
these may be unaware of cv. Salamain. Many informants 
can provide names and some kind of information on four 
or five different borrucheras. The ‘‘average’’ landowner 
may own several ‘Buyés’ trees but probably has no eco- 
nomically important cultivar. Both his knowledge and 
his curiosity extend beyond the Jorracheras in his own 
garden. Following is a summary of economic uses. 


1. ‘Guamuco’—A poultice of the flowers, together with 
‘Culebra’ leaves and the stems and leaves of Phenawx 
integrifolius Wedd. is used to treat rheumatism. The 
leaves are sometimes heated and tied over swollen 
infections, or an infusion of the leaves may be used 
to bathe infections. 


2. ‘Sangre’-—Same as ‘Guamuco’. 


3. ‘Buyés’—Leaves of this cultivar are sometimes pow- 
dered and applied topically with other drugs to relieve 
rheumatic pain. 


One informant said he always uses the fruit of this at 
maize planting time. He plants a few fruits along with 
the maize in order to prevent grubs or other soil pests 
from eating the germinating seed. This interesting ges- 
ture to protect the crop can scarcely be a general prac- 
tice, however, because fruiting is rare. This is the only 
use ever encountered for any Datura fruit or seed in the 
Valley of Sibundoy. 


4. *‘Dientes’—Same as ‘Buyés’. 


5. ‘Ocre’—No use encountered. 
6. ‘Biangin’—The leaves and flowers can be ground 


[ 187 ] 


finely and mixed into a dog’s food prior to a hunting 
expedition, supposedly to make the animal locate 
more game. ‘The behavior of a dog so intoxicated 
would be most interesting to observe, but hunting is 
rarely practiced in the Valley today. 


An herbarium specimen from Sibundoy (Idrobo 2231; 


COL) records this note on biangdn borrachero: ‘Taken 
as a stimulant when feeling fatigue and also as an hallu- 
cinogen.”” I did not learn of these uses by the Sibundoy. 
However, one Sibundoy related how his three year old 
daughter ate part of some flowers of this near the house 
and became dizzy and incoherent, falling to the ground 
and shouting periodically for many hours thereafter. 


Yepes (88) included this in his account of intoxicating 


plants used by a medicine-man from Santiago (not of 
the Sibundoy tribe). 


lw 


9. 


ls 


*“Amaron’—The leaves of this cultivar are sometimes 
used as a suppurant and as an anti-rheumatic. 


‘Salaman’—This cultivar is the rarest, being known 
only in one garden, and its owner considers it the most 
toxic of all the Daturas. He uses the leaves, along 
with those of ‘Quinde’ and ‘Culebra’ in an infusion for 
bathing rheumatic limbs and joints. He has also used 
both the leaves and flowers for their psychotropic 
effect, as described below. 


‘Quinde’—This is the most widely known of the eco- 
nomically important cultivars. An infusion of the 
leaves is employed both as an anti-rheumatic and as 
a vermifuge, and the leaves are applied topically as a 
suppurant. The leaves and occasionally the flowers 
are used psychotropically, probably more often than 
any other Sibundoy Datura (cf. below). 


[ 188 | 


10. ‘Munchira’—The minute and highly toxic leaves are 
employed medicinally as an anti-rheumatic, emetic, 
carminative, vermifuge and to treat erysipelas. Their 
psychotropic use is described below. 


11. ‘Culebra’—This cultivar has received more attention 
from botanists because of its especially curious mor- 
phology, and because of its description by Schultes 
in 1955, asa distinct genus, Methysticodendron. De- 
spite the numerous reports on its medicinal, and 
especially psychotropic, importance, my investiga- 
tion showed it to be less important to the present 
day Sibundoy than either ‘Quinde’ or ‘Munchira’. 
I found it used against resfrio (‘colds,) and against 
erysipelas, and more often than other cultivars, 
against rheumatism. 


Schultes reported the use of culebra borrachera in 
treating swollen joints, combating chills and fevers, and 
in divination, prophecy, therapy, and learning ‘‘witch- 
eraft’’ (29). 

Theilkuhl, who visited the Sibundoy in 1956, was also 
able to learn of the importance of this plant as a psycho- 
tropic drug, as well as medicinally in the form of poul- 
tices, ‘‘to combat rheumatism, swollen infections, muscle 
cramps, erysipelas, and in the form of baths, colds’’ (35). 


12. ‘Andrés’—One informant said he had used the flow- 
ers psychotropically. 


Psychotropic use 


Literature dealing with the Daturas of the Valley of 
Sibundoy has emphasized their importance as psycho- 
tropic, or mind affecting, drugs (27, 29, 80, 31, 82, 38, 
36, 38). Writings based directly upon experience among 
the Sibundoy are those of Schultes (1955) and Theilkuhl 


[ 189 ] 


(unpublished thesis, 1957) dealing only with ‘Culebra’, 
and of Pedro Juajibioy (1960, MS.). Yepes interviewed 
Inga-speaking medicine-men travelling through Popayan 
to the north (88). 

Because of the possibility that the Sibundoy learned 
the use of some of the Datura cultivars from the nearby 
Inga-speaking peoples of Santiago, Col6n and San An- 
drés, it is well to include here the following information 
collected from some of their medicine-men (88): 


Borracheras. (Daturas). They are characterized as ‘calientes (1) 
equal to fire.” They distinguish more toxic and less toxic species, 
and give preference to the latter for divinatory purposes. When 
they take them, they have an assistant accompany them because 
they produce furious intoxication beyond certain dosages. 
Borrachera. ‘Lengua de Tigre or Tinye’ (2). In small quantity, it 
gives strength. Beginning by cooking three pairs of leaves or one 
flower, one progresses to larger dosages with time, and in this form 
it serves to make divination, diagnosis and witchcraft | maleficio]. 
Biangadn borrachera. During the wane of the moon, 2 pairs of leaves 
are given to the dogs so that they will go out as hunters (3). 
Danta borrachera. It is given to hunting dogs so that they may 
follow easily the trail of any animal (3). 


(1) Calientes (‘hot’ plur.), medicines which ‘“‘cure the diseases... 
produced by cold.’’ (38). 
(2) Identity unknown, but apparently referable to D. candida Saff. 
(3) Apparently D. candida ‘Biangan’. 
The Sibundoy [Inga-speaking native of the Valley of Sibundoy, 


not a Sibundoy | knows intoxication with borrachero by the dryness 
of the throat and the dilatation of the pupil (38). 


Schultes, who investigated the plants among the Si- 
bundoy in 1942, 1946 and 19538, summarized his findings 
on the psychotropic aspects of culebra borrachera in these 
words (29): 

This intoxication, resorted to by the witch-doctors only for very 

important or difficult cases of divination, prophecy or therapy, usu- 


ally lasts for two full days and sometimes may persist for four — 
with a long period of complete lack of consciousness. Missionaries 


[ 190 | 


who have been at work in the Valley of Sibundoy for more than a 
quarter of a century suspect that the death of one aged witch- 
doctor may have been due to an overdraught of this narcotic. 
Smaller doses of the drug are administered to boys who are study- 
ing witchcraft. Certain of the medicine-men’s secrets apparently 
are imparted only when the novitiate is “‘under the protection’’ of 
this narcotic. 


Both Schultes’ and Theilkuhl’s descriptions of the 
method of preparation of the drug are nearly the same 
as that described below for the other cultivars. 

Theilkuhl, after visiting the Sibundoy in 1956, wrote 
of this clever use of eulebra borrachera (85): 

. it is suspected that occasionally unscrupulous Indians use it 
as a ‘burundanga’, that is, to deprive outsiders of their senses and 
rob them. 

It seems that the ‘médicos’ or ‘curacas’ take an aqueous macera- 

tion of the leaves to produce hallucinations, during which they say 

they see the solution of difficult cases of divination, prophecy or 
diagnosis. One of the “médicos’ questioned affirms that the ‘cule- 
bra borrachera’ is employed in a manner similar to that of ‘yagé’ 

[ Banisteriopsis spp.; (6)], the focal point of Amazonian medical 

practice. 


Yepes, Schultes and Theilkuhl use the words divina- 
tion, prophecy, diagnosis and witchcraft in describing 
the psychotropic use of Datura, yet no Sibundoy I en- 
countered stated such reasons for using any Datura drug. 
Nor has Haydée Seijas found these usages during more 
than one year of investigating Sibundoy ethnomedicine 
(pers. comm.). Most of those with whom I talked ap- 
peared to consider the drugs hallucinogens, but I failed to 
record precisely their statements. Had this been done by 
myself and the other observers, we might assess the psy- 
chological, as well as social, significance more accurately. 

The Sibundoy observer, Pedro Juajibioy, has recorded 
five instances of intoxication with Datura among his 
acquaintances (20). In three cases, it is stated that the 
leaves were taken to cure disease: agua blanca (‘white 


[ 191 ] 


water’) and stomach ache. A gua blanca reters probably 
to gonorrhea (Seijas, pers. comm.). Only in one case 
does he mention the identity of the cultivar—‘Munchira’. 
The following experience may be due to use of D. can- 
dida ‘Culebra’: 
The first time I drank six leaves at night. I got drunk. I saw woods 
full of trees, people from somewhere else, animals, stumps, pastures 
full of all kinds of snakes that came up the slopes of the pasture 
all in green. ‘They coiled to bite me. As the intoxication took 
greater hold of me, the house started to revolve against the world, 


and the same with the things in the house, ete. But the snakes 
stretched for the kill! 


Such horrible hallucinations may be commonplace. 
The Sibundoy appear to have an instinctive fear of snakes, 
despite the supposed absence of poisonous ones in the 
Valley. One medicine-man who had never taken ‘Cule- 
bra’ thought that if he did he would see ‘‘ugly things, 
snakes’. 

Juajibioy (ibid.) gives another example which was 
surely entertaining for observers: 

I took . . . only six leaves. I got drunk after an hour [and?] my 

vision went dim. During the day I saw unknown people duplicated 

—that is, one person in twomen. I felt crazy. I started running, 

then took off my clothes, and ran around naked in the garden, and 

showered myself with the piles of dirty weeds left by the hired 
men who were working at that moment. I also insulted them freely. 

And I went about kissing several tree trunks thinking they were 


my fiancée. Later I took to writing letters. I went out to the pasture 
with a rope to catch a horse to ride, but it turned out to be a dog. 


Despite these reports, I did not succeed in locating 
any native of the Valley who had employed D. candida 
‘Culebra’ for its psychotropic action. One prominent 
medicine-man has in his garden eight plants of this clone, 
perhaps one quarter of all the plants extant in the Val- 
ley. While he regularly takes the psychotropic Banis- 
teriopsis (6) and has tried several of the Daturas, he has 
not taken ‘Culebra’ and knows of no one who has done 


[ 192 


so. Frequently natives do not specify which borrachera 
they refer to unless questioned, and outsiders may easily 
assume culebra is intended. 

Seijas reports that borrachera is sometimes added to 
chicha, the nutritious daily beverage of low alcohol con- 
tent (pers. comm.). However, she adds that many con- 
sider this an antisocial practice, since it often results in 
disorderly behavior and fighting. 

A dozen cases of intentional intoxication with the 
other three important Datura cultivars have come to 
my attention. 

As psychotropic drugs, it is “Quinde’, ‘Munchira’ and 
(reportedly) ‘Culebra’ which are employed principally. 
The juice of the leaves, or occasionally of the flowers, is 
taken, usually alone and unheated. Depending on the 
size of the leaves, between one leaf and twenty-four 
leaves (‘‘twelve pairs’’) are taken. Normally the leaves 
are taken in pairs, and sometimes (Seijas, pers. comm. ) 
only as even numbers of pairs. While one leaf is sufh- 
cient in the case of the large-leaved ‘Salaman’, two or 
four of the smaller ‘Quinde’ leaves are used. Between 
eight and twenty-four of the minute *“Munchira’ leaves 
are required. The leaves are usually macerated and 
crushed and the expressed juices taken alone or some- 
times mixed with a jigger of water or trago (alcoholic 
distillate), partly for convenience. Sugar is sometimes 
added to the juice to make it more palatable. One in- 
formant heated ‘Munchira’ leaves ina very small amount 
of water to extract the juices. 

One medicine-man made the drug by expressing the 
juice from one flower each of ‘Salaman’, ‘Quinde’ and 
‘Munchira’, thus obtaining about 15 cc. to which he 
added sugar. Another individual used only two flowers, 
both of ‘Quinde’. 

The psychotropic use of these Daturas is not restricted 


[ 193 | 


Salvador Chindoy and his wife posing beside his aged “Munchira’ 


tree. (A prop prevents the crown from leaning on the roof of the 


house. ) 


to medicine-men; they are in exclusive possession of 
neither the trees nor of the knowledge of how to prepare 
the drug. However, it is likely that in many cases a 
medicine-man prepares or directs the preparation of the 
drug, especially with regard to dosage. 

The descriptions of the psychotropic effects, related 
to me by those who have experienced them, are in re- 
markable agreement. It stuns them, makes them lose 
their senses, and leaves their mouths and throats so un- 
pleasantly dry that they can neither swallow nor talk for 
many hours. Their vision is disturbed, bringing objects 
nearer and farther away, and some experience visual hal- 
lucinations, especially of animals that they fear. One 
woman said that taking ‘Munchira’ made her feel 
stronger and younger. Unfortunately, the dominant 
memory is of the long hours with a dry, constricted 
throat which prevented eating, drinking and speaking. 
Informants discussing Datura narcosis often continue on 
with their experiences with biaxti (Banisteriopsis spp.), 
experiences that they usually recall as more pleasant and 
more valuable (6). 


Chemistry 


Chemical analyses of Datura candida ‘Culebra’ 
(Methysticodendron Amesianum) were done within five 
years after its description in 1955 by Schultes. 

Theilkuhl found hyoscyamine and atropine in the 
leaves, though he believed the last-named was produced 
probably during the isolation process (85). Pachter and 
Hopkinson, who worked with thirteen kilograms of dried 
leaves and stems, found four alkaloids present, of which 
{-scopolamine accounted for 80 per cent (25). Atropine 
was also present, but two others in smaller amount were 
not identified. The plant materials for both studies were 
obtained in the Valley of Sibundoy or originated from 


[ 195 ] 


such material. ''o my knowledge, no other Datura ma- 
terial deriving from populations in the Valley has been 
analyzed chemically or pharmacologically. 

The alkaloids of the genus Datura have been subjected 
to intensive study. For a concise review by Edward 
Leete see Avery et al. (1). The principal alkaloids of the 
tree Daturas are the tropanes hyoscine (scopolamine), 
hyoscyamine and atropine. W.C. Evans and his co- 
workers are actively investigating these and closely re- 
lated alkaloids in Datura cornigera (white flowered) and 
D. sanguinea (11, 12, 18, 14). Their efforts promise to 
demonstrate alkaloidal composition and biogenesis in all 
the tree Daturas. They are currently analyzing several 
of the cultivars described herein. The roots of tree Da- 
turas, like those of the herbaceous forms, contain in ad- 
dition to the above named alkaloids, esters of tighic acid 
with 3a-hydroxy-, 8a, 6 B-dihydroxy- and 8a, 6 6, 7 B- 
trihydroxytropane. Ditigloyl esters appear to be present 
in all the Datura species examined. D. sanguinea con- 
tains acetoxy derivatives, one occurring as the mixed 
ester 8a-tigloyl-6 B-acetoxytropane. 


a) / ‘ . 
Pharmacology 


The psychotropic effects of the crude drugs upon the 
Sibundoy have been noted above. 

Atropine, hyoscyamine and scopolamine are of con- 
siderable importance in modern medical practice, but the 
commercial sources of the drugs at present are other 
solanaceous genera, Duboisia and Hyoscyamus. 

Henry summarizes the pharmacologic effects of these 
as follows (17): 

When administered internally in toxic doses, atropine at first stimu- 

lates but eventually depresses the central nervous system, giving rise 

to hallucinations, incoherent speech, delirium and convulsions, 


followed by stupor and coma, It paralyzes muscles and secretory 
glands to the effects of stimulation by post ganglionic, cholinergic, 


[ 196 | 


nerve fibres. It is to this action that the dryness of throat and 
mouth characteristic of belladonna poisoning is due. 

The natural alkaloid, 7-hyosecyamine and its d-isomeride resemble 
atropine (d/-hyoscyamine) qualitatively in action. . . . 

{ Hyoscine (scopolamine) | has an action similar to, but more tran- 
sitory than, that of atropine on the peripheral, cholergic, autonomic 
nervous system. Its action on the central nervous system is differ- 
ent. Generally it induces a feeling of fatigue and drowsiness pass- 
ing into sleep. In some cases there may be a preliminary stage of 
excitement, and with large doses excitement indistinguishable from 
that of atropine intoxication may occur. The respiratory centre is 
depressed from the start. The /-isomeride has the more powerful 
peripheral action, although the central action of both isomerides is 
the same (p. 106-7). 


Claus and Tyler (10) add that atropine applied locally 
causes a ‘“‘dullness or slight paralysis of the sensory 
nerves’, thus easing pain. This action probably relates 
to the frequent use of leaves in treating rheumatism. 

General statements such as these give a good indica- 
tion of the types of pharmacologic activity which proba- 
bly will be demonstrated in the tree Daturas, and suggest 
the basis for both their medicinal and psychotropic use 
by the Sibundoy. However, because the Sibundoy Da- 
turas are constant hosts to several viruses which may 
affect the cultivars differently, and because the Sibundoy 
have encountered greatly differing levels of toxicity, it 
seems very probable that an eventual understanding 
both of the biogenesis and of the pharmacologic activity 
of the Sibundoy Datura alkaloids will constitute a unique 
chapter in our increasing knowledge of the solanaceous 
tropane alkaloids. 


Key to Datura in the Valley of Sibundoy 


A. Exposed corolla mostly red, leaves soft-pubescent D. sanguinea 

a. Corolla tube yellow at the middle . . . . . cv. Guamuco 

aa. Corolla tube green or red at the middle, lacking 
Wellow entirely Ǥ < & 4s & # & @ ss 

AA. Corolla lacking red, leaves not soft-pubescent . . D2. candida 


[ 197 ] 


ev. Sangre 


Key to Datura (cont. ) 
a. Corolla yellow-orange. . .......... . ev. Ocre 
aa. Corolla white 
b. Corolla very deeply lobed, filaments apically 
contorted, leaves long-linear. . . ev. Culebra 
bb. Corolla slightly lobed, filaments destehe. edee: 
not long-linear 
c. Leaves with regular venation and uniform margins 


d. Leaves dentate .......... .. . ev. Dientes 
dd. Leaves entire 
Pistil exceeding 30 cm., corolla 31-43 em. ev. Andrés 
Pistil less than 25 cm., corolla 20-30 cm, 
Leaves yellowish, pistil 22-24 em. . . . cv. Biangan 
Leaves not yellowish, pistil 14-21 em. . . cv. Buyés 
ec. Leaves with irregular venation and variously 
erose margins (cf. Plate LII) . . . . . . evs. Amarén 
Salamin 
Quinde 
Munchira 


Taxonomy of the genus Datura 
in the Valley of Sibundoy 


1. Datura sanguinea Ruiz & Pavon F). Peruv. 2 
15. 1799. 

Brugmansia bicolor Persoon Syn. Pl. 1: 216. 1805; 

Lindley Bot. Reg. 20: pl. 1789. 1884. 

B. sanguinea 1). Don in Sweet Brit. Fl. Gard. II. 3: 

pl. 272. 1885; Lagerheim Bot. Jahrb. Engl. 20: 662. 

1895. 

Small trees, 2-5 meters in height; leaves soft- 
pubescent, the lower repand to angular, the upper usu- 
ally entire; flowers 17-25 cm. long, the corolla with a 
short-toothed, recurved to strongly reflexed (rarely erect) 
limb, the tube typically green within the calyx, yellow 
at the middle and red or orange-red (rarely yellow- 
orange to bronze) toward the limb, the ribs yellow; an- 
thers 12-18 mm.; stigma obtuse; fruit ovoid (rarely 
oblong-elliptic), the base rounded to acute (rarely 
acuminate) and mucronate, in cross section usually bi- 


[ 198 | 


PLaTE LIV 


aaaemaaes ——— “ eee ee es eee aes 


Datura sanguinea 
y R.& P. | 


cv. GUAMUCO 


“NS 


| S. Gnic- es 
| Peek Re | | 
aecinimee se ee erie eater Ay | eee —= 


| so Fa 
{ 
| 


Datura sanguinea R. & P. cv. Guamuco. Flowering and fruiting 
branches, “<«. 


~ 


laterally flattened, or 4~5-sided to nearly round with 2 
or 4 longitudinal sulci: seeds verruculose and lacking ¢ 


_ 


greatly thickened testa. 


la. ‘Guamuco’ n. cv. Plate LIV. 

Calyx broad, 1 8-lobed, 6.7-12.2 em. long; corolla 
17.6-21.2 cm., yellow at the middle, red toward the 
limb; stamens 15.7-19.2 em. (including their adnation 
to corolla), anthers 15-18 mm. ; pistil 15.5-19 em. : fruit 
ovoid to conical, bisulcate, with half-persistent calyx, 
rare. N=12 (Bristol 1114). 

The pollen very closely resembles that of D. candida. 
Kxamination of 500 microspores of one plant showed 
58.8 per cent to be abnormal. 

Common names: Guamuco, Guamucu borrachera. 

Guamuco is anon-Kamsa term, also used occasionally 
for Spigelia pedunculata A BK. It may relate to euambia 
‘poison’, for the Sibundoy know both plants to be highly 
poisonous, 


CotomBia: Comisaria del Putumayo: Valle de Sibundoy, alt. ca. 
2200 m.—1.5 km. S Sibundoy. “Tree 3 m.: corolla yellow below, 
reddish orange above, veins yellow above; anthers white. In cultiva- 
tion: very rare.”’ 20-XII-1962, 479* (ECON), —San Andrés. ***“Gua- 
muco’. Shrub 2.5 m.; corolla orange-red, limb recurved. Cultivated, 
infreq.’” 18-IIT-1963, 65.7. (COL, ECON, PASTO, US).—3-5 kms. 
S Sibundoy. ** ‘Guamuca borrachera’. Tree 3.5 m.; a few lvs. en- 
tire; corolla tube green, becoming yellow; throat and limb red, be- 
coming orange during anthesis; anthers white, pollen abundant: no 
fruit. Indian garden, very infreq.’’ 12-VI-1963, ///4 (COL, ECON, 
BISH, PASTO,S, US). —2 kms. SE Sibundoy. **‘Guamuco’. Shrub 
2m.; corolla tube green, becoming light orange, limb and throat 
deep red, becoming lighter. Indian houseyard, very infreq.”* 27- 
VILI-1968, 7345 (COL).—Colén. ***‘Guamuca borrachera’. Tree 4 
m., trunk diam. 23 em., young bark brown: calyx slightly purplish ; 
corolla red-orange except greenish yellow where emerging from calyx, 
and the ribs yellow, limb recurved; fr. ovoid or triangular, acuminate 
with 2 furrows, the calyx half-persistent ; seeds dark brown,.’’ 23-IX- 
19638, 1420 (COL, ECON, BISH, kK, PASTO, US).—Sibundoy. 29- 
V-1946, Schultes & Villarreal 7689 (COL, ECON, US). 


* Collection numbers are those of the author, unless otherwise specified. 


{ 200 | 


1b. cv. Sangre. 

This cultivar, described in Bot. Mus. Leafl. Harvard 
Univ. 21: 236 (1966), is distinguished by the corolla 
color, which is deep red (basally green), entirely lacking 
in yellow. The flower size and shape is variable; in the 
Valley of Sibundoy, the calyx is narrow, terminating in 
a single apicule, 8.8-9.5 em. long, the corolla 17.6—21 
em. long. Stamens (incl. their adnation to corolla) 15.4— 
18.9 cm., anthers 13-14 mm. and pistil 16.9-19 cm. The 
oblong, bisuleate fruit has a persistent calyx. 

Common names: Guamuco, Guamuco borrachera, as 
for the first cultivar. 

Cotompra: Comisaria del Putumayo, Valle de Sibundoy, alt. ca. 
2200 m.—San Andrés. ‘‘*Guamuco’. Shrub 2.5 m. ; corolla red, limb 
spread somewhat, but not rolled upward as in 653. Cultivated.’’ 18- 
Il1-19638, 652 (COL, ECON, PASTO, US).—San Andrés. “‘Arbores- 
cent 2.5 m. ; exposed corolla red (no yellow present); fr. 9055 mm., 
ovoid, 5-ridged, not flattened. Roadside, common.’’ 20-VIII-1963, 
1309 (COL, ECON, US). 

2. Datura candida (Pers.) Safford in Journ. Wash. 
Acad. Sci. 11: 182. 1921. 

D. arborea Ruiz & Pavén Fl. Peruv. 2: 15. pl. 128. 

1799; non D. arborea L. 

Brugmansia candida Persoon Syn. Pl. 1: 216. 1805. 

D. aurea Lagerheim Gartenfl. 42: 38. 1893; Safford 

ibid. 186. 1921. 

B. aurea Lagerheim in Bot. Jahrb. Engl. 20: 664. 

1895. 

B. arborea Lagerheim ibid. 663. 

D. affinis Safford ibid. 186. 

D. Pittiert Safford tbid. 187. 

Small trees, 3-5 meters in height; leaves glabrous or 
slightly pubescent, ovate or oblong-elliptic, entire or 
coarsely dentate; calyx 1.5-8 cm. broad, 1—4-toothed; 
the slender basal part of the white corolla wholly en- 
closed by the calyx, the limb flaring broadly with long 


[ 201 ] 


(4-9 em.) recurved teeth; anthers distinct; fruit oblong- 
cylindric to fusiform and lacking a persistent calyx; seeds 
angular, with a greatly thickened, suberose testa. 


2a. ‘Buyeés’ n. cv. Plates XLEX and LV. 

Ratio of leaf width to length .846—.577, calyx 8.5-16.1 
em., corolla 19.6—-29.3 cm., stamens (incl. adnation of 
filaments to corolla) 18.2-19.9 em., anthers 80-41 mm., 
pistil 14,8-21.2 em. N=12 (Bristol 1117, 1266). 

This is the common borrachera found throughout the 
Valley in inhabited locations and where there is evidence 
of sites of former habitation. Cv. Buyés is the most 
variable of all the D. candida cultivars. Several to many 
clones are included here, but they are only obscurely 
differentiated, and neither the natives nor the people of 
Spanish descent distinguish among them. Differences 
in the splitting of the calyx, diameter of corolla tube, 
length of corolla and length of corolla teeth, as well as 
others, can be seen. 

Common names: Borrachera, borracherushe, buyés 
borrachera, buyés borracherushe, borrachera de agua, 
foripundo, floripondio blanco, guamuco blanco, guamuco 
Horipundo. 


Borrachera ‘inebriant’ is from the Spanish borracha 
‘wine skin’, whence borracho ‘drunkard’. The Sibun- 
doy believe it to be a word of their own language, 
Kamsa, an indication of its long usage among them. 
Elsewhere in Colombia J. candida is known as borra- 
chero ‘inebriating tree’. 

Borracherushe is a variant, and more typically Kamsa 
form, of the first. 


Buyés ‘water’ is of unexplained application here, but 
perhaps it refers to the plant’s preference for a wet 
site, and the common habit of planting it near ditches. 


[ 202 ] 


PLatTE LV 


yo Se 
y Datura candida 
— CPERS.) SAFF. | 
cv. BUYES | 


atti pense 


nee eal 
Me tiga”, 


we coe MUI Ce ee eerie ys 


Datura candida (Pers.) Saff. ev. Buyés. Flowering and fruiting 


branches, 4. 


Floripondio (Spanish) ‘floribunda’ refers to ‘‘the size and 
abundance of its flowers”” (88). 


A gua (Spanish) ‘water’, equivalent to buyés. 
Blanco (Spanish) ‘white’, for the flower color. 


Guamuco (language?) is the name in the Valley and else- 
where for the red-flowered D. sanguinea. 


Buyés borrachera and its variants are employed with 
about equal frequency by the Sibundoy, while the re- 
maining names are used only by immigrants from Narino 
and elsewhere in Colombia. 


CotomsiA: Comisaria del Putumayo, Valle de Sibundoy, alt. ca. 
2200 m.—Sibundoy, collados al norte del pueblo. “‘Nombre castel- 
lano: ‘borrachera’. Large bush. Fl. white.’’ 12-II-1942, Schultes 
3207 (GH, NY).—Sibundoy. ‘“* “Bui-ish borrachera’. Tree 4 m.; 
corolla white, fragrant, limb cordate between teeth. Calyx teeth 2-3. 
Borders, freq.’’ 26-I1V-1963, 887 (COL, ECON, BISH, K, PASTO, 
S, US).—Sibundoy. ‘‘‘Bui-ish borrachera’. Arborescent 2.5 m.; 
corolla white. Borders, frequent. ’’ 26-IV-1963, 889 (ECON).—8 km. 
SW Sibundoy. ‘‘Tree 4 m.; corolla white. Infreq. in borders.’’ 9- 
V-1968, 999 (COL, ECON, PASTO, US).—4 km. SW Sibundoy. 
‘**Bui-ish borrachera’. Tree 4m. Secondary woodland.’’ $-VI-1963, 
1098 (COL, ECON, K, PASTO, US).—Sibundoy. ** ‘Bui-ish borra- 
chera’. Tree 3.5 m.; corolla white; frs. green, few. Hedge row, 
frequent.’” 10-VI-1963, 7109 (COL, ECON, US).—3-—5 km. S Sibun- 
doy. ** ‘Bui-ish borrachera’. Tree 5 m.; corolla white; fr. green. 
Borders, common.”’ 12-V1-1963, 7117 (COL, ECON, US).—Sibun- 
doy. ‘‘‘Bui-ish borrachera’. Tree 4 m.; firs. pendulous, corolla 
white: fr. green, indehiscent. Border, frequent.’’ 10-VI-1963, 1/93 
(COL, ECON, BISH, K, PASTO, S, US).—Sibundoy. ** *Bui-ish 
borrachera’. Tree 4 m.; corolla white, limb recurved. Borders, fre- 
quent.’’ 22-VII-1968, 7255 (COL, ECON, BISH, K, PASTO, §S, 
US).—Colén. ‘‘Tree 2.5 m.; corolla white. Border, Indian garden. ”’ 
25-VII-1963, 1266 (COL, ECON, BISH, K, PASTO, S, US). 
km. SW Sibundoy. “*“*Bui-ish borrachera’. Arborescent 2-5 m. ; firs. 
pendulous, corolla emerging pale yellow, white when fully open; 
anthers white; stigma very pale yellow. Borders, common.’’ 19-IX- 
19638, 1388 (ECON). 


9 


2b. ‘Dientes’ n. ev. 
Leaves dentate, ratio of width to length .550, calyx 


[ 204 | 


11.1-13.4 cm., corolla 22.5-25.7 cm., stamens (incl. 
adnation of filaments to corolla) 14.3-16.5 cm., anthers 
32-88 mm., pistil 15.8-17.8 cm., fruiting occasionally. 

The distinguishing feature of this cultivar is the den- 
tate leaf’ margin which is usually restricted to the distal 
half of the leaf, but which may occur throughout the 
length. Almost all of the plants in the Valley of Sibun- 
doy occur in a single locality two to three kilometers 
southeast of the town of Sibundoy. Along three main 
trails covering two kilometers here, the many landowners 
have planted nearly 250 of these trees. 

Common names: Buyés borrachera, ete., the same as 
for D. candida ‘Buyés’. 
CotompBia: Comisaria del Putumayo, Valle de Sibundoy, alt. ca. 
00 m.—-2 km. SW Sibundoy. “*Trailside, very infreq.’’ 14-I1V-1963, 
4 (ECON).—8 km. S Sibundoy. “*‘Bui-ish borrachera’. Tree 4 m. ; 
corolla white; fr. green, pericarp drying and decaying but indehis- 
cent. Border, infreq.’’ 15-VI-1963, 1/22(ECON).—2.5 km. S Sibun- 


eee 


doy. Bui-ish borrachera’. Tree 4 m.; corolla white, pendulous; 
frs. broadly fusiform, indehiscent. Borders.’’ 2-X-1963, 1435 (COL, 
ECON).—2.5 km. S Sibundoy. ‘‘Arborescent 3 m.; corolla white, 
pendulous. Indian gardens, borders; frequent.’’ 2-X-1963, 148 
(ECON).—2 km. SE Sibundoy. *“Arborescent 4 m.; corolla pendu- 
lous, white, limb and teeth recurved; fr. terete, fusiform (15.5 3.5 
em. ), indehiscent. Borders, frequent.’’ 8-X-1963, 1447 (COL, ECON, 
BISH, K, PASTO, S, US). 


99 
any 
if 


Another collection (/177) has both distally dentate 
and entire leaves from the same plant. In addition, the 
calyx is very narrow with the limb partly recurved, and 
the corolla tube is also narrow. Ten to 12 trees occur in 
a single hedge row in the town of Sibundoy. During 
May and early June, 1963, the only Daturas blooming 
in the Valley were this planting and ‘Culebra’. The 
owner called this ‘‘bui-ish borracherushe’’ and said that 
the first cuttings had been brought here from further 
south in the Valley, where most of the Sibundoy reside. 


[ 205 ] 


CoLtomsia: Comisaria del Putumayo, Valle de Sibundoy, alt. ca. 
2200 m.—Sibundoy. “* ‘Bui-ish borracherushe’. Tree 3.5 m.: lvs. 
entire or distally toothed; calyx slender, limb curling; corolla white. 


Hedge row, infreq.’” 11-VI-1963, 77177 (COL, ECON, BISH, K, 
PASTO, S, US). 


2c. ‘Ocre’ n. cv. Plate LVII. 

Leaves entire, the base acute, ratio of width to length 
.402~-. 498, calyx relatively narrow, 12.8-15.9 cm., corolla 
long, 25.4-81 em., ochre color, stamens (including their 
adnation to corolla) 16.4-22.6 cm., anthers very long, 
40-44 mm., pistil 17.8-23.8 em. Fruit not seen. N=12 
(Bristol 1267). 

This very rare cultivar is unique in the light orange or 
ochraceous color of its flowers. ‘wo plantings are known, 
a single tree near Sibundoy, and a short hedge row in 
Coloén. The present owners of these trees with strikingly 
different flowers showed little interest in them. 

Common names: The only name given this cultivar 
by its owners is borrachera (‘inebriant’) in Sibundoy and 
borrachero (‘inebriant tree’) in Colén. In Sibundoy, the 
response tsushie borrachera (‘yellow inebriant’) was even- 
tually evoked, but it was clear that tswshie was a purely 
descriptive word and not an established phytonym. 

CotomBia: Comisaria del Putumayo, Valle de Sibundoy, alt. ca. 
2200 m.—Coldén. “‘ ‘Borrachero’. Tree 4.5 m.; corolla light orange; 
not fruiting. Indian garden, very infreq.’’ 25-VII-1963, 1267 (COL, 
ECON, BISH, K, PASTO, S, US).—1.5 km. SW Sibundoy. ‘‘‘Tsa- 


shie borrachera’. Tree 3 m.; corolla light orange. Indian garden, 


very infreq.’’ 12-VIII-1963, 7297 (COL, ECON, PASTO, US). 


2d. ‘Biangan’ n. cv. Plate LVI. 

Leaves yellowish, entire, ratio of width to length, 
.818-.488, calyx length 10.2-16.3 cm., corolla long, 
28-32.9 cm., in some the slender basal part of the tube 
well exposed, stamens (including their adnation to co- 
rolla) 19.9-23.2 cm., anthers 837-43 mm., pistil 21.6-24.4 


[ 206 ] 


PiaTeE LVI 


pica acim ine, 


Seat AON, _ Datura cantida 


NG CPERS.) SAFF. 
SS) cv. BIANGAN 
f. SAY : ( ? 


4 


YY 
Cs 


L See i 


Datura candida (Pers. ) Saf, cv. pinaaia: Flowering ae fruiting 
branches, +4. a, Cross sections of two fruits from the same tree, 4. 


em., fruit oceasional, narrowly fusiform, somewhat 
grooved and ridged, 2 or 8 locular, calyx persistent or 
ecaducous. N = 12 (Bristol 1431). 

Cy. Biangan is easily recognized by the yellowish 
foliage and, if they are available, the unusual fruits, re- 
sembling those described by Safford (28) for D. swaveo- 
lens. A photograph of ‘Biangin’ flowers appears in Perry 
(27), page 335, centre figure. 

Common names: Biangdn borrachera, chontaruco bor- 
rachera, danta borrachera. 

The Sibundoy know two deer, a small one (biangun), 
said to be only one half meter high, which lives near the 
marsh in the centre of the Valley, and a larger one (mon- 
gqjo) of the high mountains and paramos. Chontaruco 
(‘biangiin’) may be an Inga term, while danta (‘tapir’) 
is a widespread indigenous word adopted by Spanish. 
The Sibundoy think of D. candida ‘Biangin’ as a stimu- 
lant for dogs on hunting trips, but whether their names 
refer to inebriated dogs capturing game animals or to 
the possibility of game animals becoming inebriated 
themselves by browsing on the leaves, is unclear to me. 


Cotompra: Comisaria del Putumayo, Valle de Sibundoy, alt. ca. 
2200 m.—Sibundoy. ‘* ‘Danta borrachera’. Bush’’. 29-V-1946, 
Schultes & Villarreal 7638(US).—Sibundoy. *“N.v.Danta borrachero,’ 
*Biangan borrachero’ (Kamsa). Arbusto de 3m. Hojas amarillentas. 
Flores vivas, no vistas. (Se toma como estimulante cuando se siente 
cansancio y tambien como alucinégeno).’” 12-VI-1956, Idrobo 2231 
(COL).—8 km. SE Sibundoy. ‘‘ “Bui-ish borrachera’. Arborescent 
3 m.; upper lvs. small, yellowish; firs. (except calyx) white through- 
out. Edge of cornfield.*’ 8-I-1963, 458 (COL, ECON, PASTO).— 
5 km. SW Sibundoy. “* ‘Biangan borrachera’, Arborescent 2.5 m., 
lvs. yellowish green, calyx becoming yellow; corolla white (between 
lobes rounded or very slightly emarginate); no fr.’’ 29-IV-1963, 
890 (COL, ECON, BISH, K, PASTO, S, US).—5 km. S Sibundoy. 
‘**Biangan borrachera’, Tree 3.5 m., lvs. yellowish; corolla white; 
fr. green (loose, persistent calyx removed), 2-3 carpellate. Border, 
very infrequent.’’ 27-VI-1963, 1162(COL, ECON, K, PASTO, US). 
—8 km. S Sibundoy. *‘ ‘Biangan borrachera’. 2 m., lvs. yellow- 
green; corolla white; frs. green, 2-3 earpellate, angular and grooved 


( 208 ] 


(2 carpellate flattened). 18-VII-1963, 7246 (COL, ECON, US).—2 
km. SE Sibundoy. ** “Biangan borrachera’. Shrub 2 m.; leaves and 
calyces yellowish; corolla white. Pasture border, infreq.”* 27-VIII- 
1963, 1348 (ECON).—1.5 km. S Sibundoy. ** “Biangan borrachera’. 
Tree 3.5 m.:; lvs. yellowish: corolla white. Indian garden, very in- 
freq.’’ 30-VIII-1963, 1356 (ECON).—1.5 km. S Sibundoy. ** “Bian- 
gan borrachera’. Tree 3 m., lvs. yellowish; firs. pendulous, corolla 
white, limb recurved but teeth hanging downward, limb from base 
of tooth to segment edge rolled backward. Indian houseyard, very in- 


freq.’? 2-X-1963, 1431 (COL, ECON, BISH, K, PASTO, US). 


Ye. ‘Amaron’ n. cv. Plate LVII. 

Leaves deformed, lanceolate to oblong, reaching 45 
cm., margin irregularly undulate and obscurely dentate, 
or not vertically undulate and strongly dentate, the 
lateral veins curving near midrib, often forming a slight 
S-curve; ratio of width to length .218-.364: calyx 9.4— 
15.5 em., corolla 20.3-27.8 em., the tube relatively 
broad, stamens 13-18.2 em. (including their adnation to 
corolla), anthers short, 28-384 mm., pistil 14-19.8 ¢m., 
fruit occasional. N=12 (Bristol 564). 

Cv. Amar6én, consisting of several clones, occurs at 
many localities in the Valley, sometimes in extensive 
hedge plantings. Probably there are in excess of 200 
trees. The variable leaves may be lanceolate to narrowly 
oblong and with the margins both horizontally and ver- 
tically undulate and sometimes distally dentate. A few 
plants have both irregular leaves and leaves similar to 
‘Buyés’ leaves in outline, but with the venation dis- 
torted. Most of the plants grow a little more vigorously 
than the other cultivars, as seen in the greater size of the 
trees, the leaves and fruits and in the thickness of young 
stems. 

Common names: 4 maron borrachera, cucu borrachera. 

A marron (Spanish) ‘boa constrictor’, a thick, heavy- 
bodied snake. Possibly the name is applied to this cul- 
tivar in recognition of the very thick, young herbaceous 


r 5 


| 209 | 


des 4 


Datura candida (Pers. ) Saff. (Upper) Foliage of ev. Amaron. (Lower) 


Imergent and open flowers of cv. Ocre. 


stems which usually distinguish this from all the other 
cultivars. 

Cucu was given by only one informant. The basic 
meaning of the word among the Sibundoy is not known 
to me. Spanish speakers use it in reference to ‘devil’ 
(Seijas, pers. comm.). Pazos (26), considering Quechua 
vocabulary in use in Colombia today, gives ‘‘Quechua, 
kuku: espantajo, fantasma’’, apparently ‘fright, ghost’. 
However, Lira (23), in his exhaustive treatment of Que- 
chua in the Inca heartland in southern Peru, gives the 
following: ‘‘K’ UKU, adj. Immaturo, no maduro, verde 
y duro, fruto verdete sin sazon. Iam. Divieso o quiste 
endurecido.’’ The familiar application divieso ‘furuncle, 
boil’ and quiste ‘cyst’ is the most likely sense of the term 
here, for this cultivar (as well as others) has been used as 
a suppurant in treating boils and cysts. 


CotomBiA: Comisaria del Putumayo, Valle de Sibundoy, alt. 2900- 
3180 m.—Road from Sibundoy to Pasto: between La Maria and Paramo 
de San Antonio. ‘“Bush 12 ft. Fl. white.’’ 1-VI-1946, Schultes & 
Villarreal 7809 (ECON, US). Valle de Sibundoy, alt. 2200 m.—3 km. 
S Sibundoy. *‘ ‘Amarr6n borrachera’. Tree 3 m.; corolla yellow upon 
emergence, white expanded. Cultigen; prop. vegetatively; infre- 
quent.’’ 27-II-1963, 564 (COL, ECON, BISH, K, PASTO, S, US). 
3 km. S Sibundoy. ““Tree 6 m.; corolla emerging yellow, white 
expanded. Cultigen; prop. vegetatively: infrequent.’ 27-I1-1968, 
565 (COL, ECON, BISH, K, PASTO, S, US).—8 km. S Sibundoy. 
“Tree 3 m.; corolla emerging yellow, white expanded. Cultigen; 
prop. vegetatively; infrequent.’’ 27-II-1963, 567 (COL, ECON, 
BISH, K, PASTO, S, US).—2 km. SW Sibundoy. ‘Tree 3.5 m.; 
corolla white. Indian garden, infreq.’’ 14-1V-1968, 775 (ECON).— 
3 km. S Sibundoy. ** “Amarr6n borrachera’’. Tree 4.5 m.; corolla 
white; frs. green, few. Indian garden, infreq.’’ 10-VI-1963, 71/0 
(ECON).—3 km. S Sibundoy. ‘‘*Amarr6n borrachera’. Tree 4 m. ; 
corolla white; frs. green, few. Indian garden, very infreq.’’ 2-VII- 
1963, 7189 (COL, ECON, US).—4 km. S Sibundoy. ** “Amarrén 
borrachera’. Tree 2.5 m.; corolla white; fr. green. Indian garden, 
very infreq.’’ 7-VII-1963, 7192 (COL, ECON, US).—Colén. ** ‘Bor- 
rachero’. Tree 3.5 m.; corolla white, pendulous; fr. green. Border, 
infreq.’’ 16-VII-1968, 7213 (COL, ECON, BISH, K, PASTO, 5S, 


US).--2 km. S Sibundoy. ‘‘‘Amarrén borrachera’. Tree 3.5 m.; 


[ 211 ] 


corolla white. Border, infreq.’’ 15-VIII-1963, 7305 (COL, ECON, 
BISH, K, PASTO, S, US).—2 km. SE Sibundoy. “‘ ‘Cucu borra- 
chera’, Tree 4 m.; corolla white. Indian houseyard, infreq.’’ 27- 
VIII-1963, 1346 (ECON). 


2f. ‘Salaman’ n. ev. 

Leaves long, the margins strongly undulate vertically, 
the distal half greatly deformed and little developed; 
calyx 9.9-15.1 em., corolla 23.3-27 cm., stamens 14.8— 
16.9 em. (including their adnation to corolla), anthers 
relatively very short, 25-28 mm., pistil 15.7-18 cm.:; 
not known to fruit. 

Cv. Salamian is perhaps the rarest of the Datura cul- 
tivars in the Valley. One Sibundoy owns three trees 
derived from a single cutting that he planted years ago; 
it was brought to him from just outside the Valley, 
where the rivers exit through a cleft in the mountains. 
That areais very sparsely settled today, and most of its 
inhabitants are not Indians. A cursory inspection of the 
area failed to locate any tree Datura. 

Common names: Salvanje borrachera (Kamsa), sala- 
manga borrachera (Inga), salaman borrachera (Spanish). 

Salamdn and its variants are employed by the same 
person, the owner of the plants, in the three languages 
which he speaks. 

CotompBra: Comisaria del Putumayo, Valle de Sibundoy, alt. 2200 
m.—Sibundoy. “*N. v. ‘Salaman-borrachero’, ‘Salvaje-borrachera’. 
Arbol de 4 m. Botones erectos u horizontales. Corolas blancas, aro- 
maticas. Los indios la distinguen como especie, pero parece ser un 
estado intermedio entre el 2207 [not seen] y 2223 [ev. Munchira].’’ 
12-VI-1956, Idrobo 2224 (COL).—1.5 km. S Sibundoy. ** “Salamaén 
borrachera’. Narcotic. Tree 5m. ; corolla white, pendulous; not fruit- 
ing. A few leaves on young, vigorous shoots are undeformed.’’ 11- 
VI-1968, 1194 (COL, ECON, BISH, K, PASTO, S, US),.—1.5 km. 
S Sibundoy. *‘ ‘Salaman borrachera’. Tree 5 m.; corolla white, pen- 
dulous; not fruiting. Indian garden, very infreq.’’ 2-X-1963, 1432 
(ECON). 


The oldest of the three trees seen is five meters high, 


(212) 


about the limit for a tree Datura, and many of its 
branches have small leaves which are only two or three 
times longer than the diminutive leaves of ‘Munchira’, 
which are characteristically the same shape. Observing 
the same trees, J.M. Idrobo had already noted the re- 
semblance to *Munchira’® (Idrobo 2224). 

Two unusual flowers (Bristol 1448) were discovered on 
one branch of a ‘Salamin’ tree. They may be interpreted 
as chimeral in origin. ‘They demonstrate the possibility 
of obtaining striking new cultivars from existing ones by 
vegetative propagation of such anomalous branches. The 
flowers are small, with all parts much shorter than nor- 
mal ‘Salamiin’ flowers, the corollas being only 21.8 and 
22.5cem. long. Most striking is the failure of the corolla 
limb to complete its growth and recurve. Instead, the 
limb flares only slightly and its margin between the 
teeth, rather than being rounded outward or cordate, is 
concave. The pistil falls far short of the stamens which 
in all other tree Daturas it exceeds. 


Cotompria: Comisaria del Putumayo, Valle de Sibundoy.—1.5 km. 
S Sibundoy. ‘* ‘Salamén borrachera’. Tree 4 m.; aberrant branch 
with 2 small firs., corolla white, limb flaring only 45 degrees, its 
edges crinkled. Indian garden, very infreq.’’ 8-X-1963, 1448 (ECON). 


2e. ‘Quinde’ n. cy. Plate LVIII. 

Leaves very irregular, frequently with two lateral 
veins and their associated laminar tissue in the basal half 
well developed, and the distal half with relatively less 
development; the flowers often somewhat smaller than 
in the other clones, calyx 10.3-15.6 cm., corolla 19.7— 
25.2 cm., stamens 13.2-15.9 cm. (including their adna- 
tion to corolla), anthers relatively short, 26-33 mm., 
pistil 18.1-18.1 cm.; fruit occasional, the calyx some- 
times persistent. N=12 (Bristol 327, 1304, 1433). 

Common names: Ngunsiana borrachera, quinde bor- 
rachera, cari borrachero. 


[ 213 ] 


Neunsiana (Kamsi) ‘hummingbird’, according to a 
Sibundoy, from the common long-tailed hummingbird 
of the Valley, the tail and wings of which are represented 
by the leaf of this cultivar. 

Quinde (Inga) ‘hummingbird’. 

Cari (Inga?’), apparently from KKH ARI (Quechua) 
‘male’ and familiarly, ‘valiant, aggressive, energetic 
person’ (23), suggesting one reason for the psychotropic 
use of this cultivar. 

Chalua borrachero, once recorded, should perhaps be 
discounted, since the Quechua term CH’ ALLU ‘ripe’ 
(23) may have been used by an [nga-speaking informant 
simply to indicate a stage of development, perhaps of 
the fruit. 

Quinde and quinde borrachera are nearly universal 
terms for this cultivar. 

Cotompia: Comisaria del Putumayo, Valle de Sibundoy, alt. 2200 
m,—Sibundoy. *‘ ‘Borrachero’’’. 25-V-1935, Garcia-Barriga 4640 
(ECON, US).—3 km. S Sibundoy. ‘* ‘Quinde borrachera’. Tree 3 
m.; corolla emerging yellow, white expanded ; bearing mature fruit. 
Cultigen; prop. vegatatively ; infrequent.’’ 27-II-19638, 566 (COL, 
ECON, BISH, K, PASTO, US).—4 km. SE Sibundoy. ‘‘ ‘Quinde 
borrachera’. ‘Tree 4 m.; corolla white; frs. green. Border; very in- 
freq.’’ 15-VI-19638, 1121 (COL, ECON, K, PASTO, S, US).—1.5 
km. SW Sibundoy. “‘Shrub 2 m.; corolla white. Indian garden, very 
infreq.’’ 12-VIII-1963, 1299 (COL, ECON, BISH, K, PASTO, S, 
US).—1.5 km. S Sibundoy. ‘‘ ‘Quinde borrachera’. Tree 2.5 m.; 
corolla white. Indian garden, very infreq.’’ 15-VIII-1968, 7304 
(ECON).—1.5 km. W Sibundoy. ‘“‘ “Quinde borrachera’. Arbores- 
cent 3.5 m.; corolla white; fr. terete, green. Indian garden, very 
infreq.’’ 22-VIII-1963, 7333 (ECON).—2.5 km. S_ Sibundoy. 
‘**Quinde borrachera’. Shrub 2.5 m.; corolla white, pendulous; 
fruit green. Indian garden, very infreq.’’ 2-X-1963, 1434 (ECON). 
—2.5 km. S Sibundoy. “‘ ‘Quinde borrachera’. Arborescent 4 m.; 
corolla white, pendulous; fr. fusiform, indehiscent. Indian garden, 
very infreq.’’ 2-X-1963, 1439 (ECON). Valle de Sibundoy, alt. 2500- 
2600 m.—2 km. NW Sibundoy. ** “Quinde borrachera’. Arborescent 
3.5 m.; corolla white. Secondary growth, far from any sign of dwell- 
ing. Solitary plant.’’ 29-V-1963, /080(COL, ECON). 3 km. N San 
Pedro. *‘ ‘Munchira’. Arbusto 2.5 m.; flor blanea.’’ 9-VIII-1963, 
Juajibioy Chindoy 190 (COL, ECON, US). 


[ 214 ] 


PiateE LVIII 


¢ \LDatura candida 
x) (PERS.) SAFF. 
“cv. QUINDE 


a 


Wf S. Lonii- 


Datura candida (Pers.) Saff. ev. Quinde. Flowering and fruiting 
branches, 4. 


ed 


Hernando Garcia- Barriga collected the earliest speci- 
men that I have seen, and I credit him as the discoverer, 
in 1935, of the peculiar tree Datura cultivars of the 
Sibundoy. 


2h. ‘Munchira’ n. cv. Plates LIIIT, LIX. 

Appearing stunted, usually 1.5-2 m. high, but reach- 
ing 38m. with age, the branches short and twisted, with 
scarcely elongated internodes, leaves relatively minute, 
variously much deformed, especially distally, calyx short, 
9.3-12 em., corolla 21.2-29 cm., stamens 15-16.7 cm. 
(including their adnation to corolla), anthers short, 27- 
30 mm., pistil 14.2-21.8 mm.; not known to fruit. 
N=12 (Bristol 1268). 

These striking, stunted plants are among the rarer, for 
there may be no more than 15 in existence. If one of 
these treelets be pollarded or cut back to the ground, 
one or more vigorous shoots arise, as with all the culti- 
vars, and there is no sign of stunting for several months. 
Perry (27) illustrated a flowering branch of this cultivar 
being examined by Salvador Chindoy and me (p. 835, 
top figure). 

Common names: Jlunchira borrachera. 

Munchira (Inga) ‘caterpillar’ alludes to the appearance 
of the leaves which seem to be (but definitely are not) 
munchiradas ‘caterpillar eaten’. 


CoLtomBia: Comisaria del Putumayo, Valle de Sibundoy, alt. 2200 
m.—Sibundoy. ‘'N. v. ‘Munchira’. Sufrutex y arbusto hasta de 2 
m.; con ramas muy tortuosas; tronco principal y ramas primarias 
cubiertas de musgo. La atrofia en las hojas se dice que prevalece en 
todos los clones. Muy conocido para tomar como estupefaciente ; vermi- 
fugo. Se dice que el principio es muy activo. Botones florales hori- 
zontales y pendulos.’’ 12-VI-1956, Idrobo 2223 (COL).—1.5 km. § 
Sibundoy. ‘**Munchira borrachera’. Narcotic. Tree 3 m.; corolla 
white; never fruiting. Indian dooryard, very infreq.’’ 26-VII-1963, 
1268 (COL, ECON, BISH, K, PASTO, S, US).—San Andrés. “‘Ar- 
borescent 1.8 m.; corolla white. Indian garden, very infreq.’* 20- 
VITI-1963, 7311 (ECON). 

[ 216 | 


PiateE LIX 


hk _ Datura candida 


CPERS.) SAFF. 
cv. MUNCHIR 


PQ EH 
ae i 
oe ol Li al Mlle = 
ft 


1. i! i Ady gs? ve ee 


ay a] itd Wnt 


(CA), 
mt ili want 


7 5 Omit 


Datura candida (Pers.) Saff. ev. } 


and habit. 


21. ‘Culebra’ n. cv. Plates LX, LXI. 
Methysticodendron Amesianum R. K. Schultes Bot. 
Mus. Leafl. Harvard Univ. 17: 2. 1955. 

Leaves narrowly linear-ligulate, ratio of width to 
length .086-.092, margin undulate: calyx distally in- 
flated, 10.8-15.8 cm. ; corolla shortest among the culti- 
vars, 15, 1—24.5 em. (excluding the relatively short teeth), 
deeply divided (through { of total length) into long spat- 
ulate lobes; stamens 12-16 em. (including their adnation 
to corolla), the filaments distally and irregularly con- 
torted, anthers 25-82 mm.; pistil shortest among the 
cultivars, 9-12.1 cm., with a variable number (2-4) of 
incompletely coherent styles exceeded by the stamens, 
the ovary often three-locular and with one or more vari- 
ably developed appendages homologous with the styles. 
Fruiting rarely or never. N =12 (3,7). (For an exhaus- 
tive and illustrated description, including details of pol- 
len morphology, see Schultes 1955. 

While strikingly different from all other Datura cul- 
tivars in the Valley of Sibundoy, there is little justifica- 
tion for excluding cv. Culebra from the genus Datura 
(3, 5, 7, 85, 87). Aberrant forms bearing a resemblance 
to this one have been observed in D. Stramonium, and 
also ina single, but very distinct, tree Datura collection 
from elsewhere in Colombia (7). 

The origin of ‘Culebra’ is unknown, but it may safely 
be assumed to derive from one of the white flowered tree 
Daturas, and it is tentatively placed with D. candida. 
With the exception of Datura ‘Andrés’, a putative hy- 
brid between D. candida and D. suaveolens Humb. & 
Bonpl. ex Willd., all of the other white flowered culti- 
vars in the Valley belong with D. candida. As with 
these cultivars, ‘Culebra’ is known only from the Valley 
of Sibundoy, and hence it can be considered a member 
of the same JD. candida complex for the present. 


[ 218 | 


PLateE LX 


— 
| 


Datura canitida 


(PERS.) SAFF. 


/ 


| 
/A 

//\ 
| 


cv. CULEBRA 


/ 
y ff 
/ 
{ / Z 
\\ ff WV 
{ ok J 
if) ™ Z 
= a os ae ey ag ee = 
= y iat Pe 
A > 
a 4 
\ 
vA 


1 


Datura candida (Pers.) Saff. ev. Culebra. Flowering branch, 95. 


Studying material grown at Bogota, Theilkuhl tound 
62.7 per cent of the flowers to be bilocular, the remainder 
being trilocular, a rare condition in the Solanaceae. With 
regard to the possible failure to form fruit, it is interest- 
ing to note that, in Bogota, the styles *‘usually begin to 
wilt before the opening of the anthers’’, thus eliminating 
the possibility of self fertilization (85). 

Schultes recalled his lost collections of fruit as fusiform 
and about six inches in length, “‘very like the fruit of 
DD). suaveolens” (29). This description, and the observa- 
tions of both 2- and 3-locular ovaries suggest the fruits 
of D. candida * Biangan’. 

The possibility ofa hybrid origin cannot be eliminated, 
but it is clear that ‘Culebra’ is not morphologically in- 
termediate between JD. candida and either 1. suaveolens 
or D. sanguinea. 

Common names: Jlutscuat borrachera, culebra borra- 
chera. 

Mutscuai (IKKamsi) and euvlebra (Spanish) mean ‘snake’, 
in reference to the long slender leaves. 

Reports (29) of other names applicable to this cultivar 
arise either from uninformed natives (to many, the trees 
are little known though a subject of curiosity), or from 
misunderstandings in widely ranging conversation (in- 
formed individuals may discuss much more than the in- 
formation solicited, perhaps without the interrogator’s 
awareness). 

Cotompia: Comisaria del Putumayo, Valle de Sibundoy, alt. 2200 
m.—Los alrededores de Sibundoy. ** Nombre kamsa :kinde-borrachera. 
Nombre castellano: culebra borrachera. Used in Sibundoy by Kamsa 
Indians as a divinatory narcotic and poison.’’ 16-I1-1942, Schultes 
3200 (COL, ECON, US). Cotype Methysticodendron Amesianum R.E, 
Schultes. )--Sibundoy. ** “Culebra borrachero’. In garden near house 
of an Indian herb doctor. Erect shrub 3-4 m. tall, fls. white, some- 
what irregular. Plant apparently sterile, as old fls. seem to drop off, 


and the people say it never sets fruit. Planted here, not seen wild.’’ 
30-11-1943, Fosberg 20406 (US).—**Tree 20 ft. tall. Cult. in Indian 


[ 220 | 


Prate LX! 


Habit of Datura candida (Pers. ) Saff. ev. Culebra. 


herb garden.”’ 21-11-1951, Villarreal 6S (ECON),.—‘'N. v. “Culebra 
borrachera’. Arbol de 2-2,50 m. de altura. Flores blaneas. Los in- 
dios mastican sus hojas para embriagarse.’” 6-I1-19538, Mora 102. 
(COL).—Sibundoy. ““Treelet 25 feet tall. Cultivated in Indian gar- 
den. ‘Culebra borrachera’. Used as a divinatory narcotic. Flowers 
white, anthers vellow.’’ 30-VI-1953, Schultes & Cabrera 20079 (GH). 
(Cotype Methysticodendron Amesianum.)—Santiago. ‘‘*Culebra borra- 
chera’. Arbolito de 2-3 m. de alto. Flores blancas. Cultivado por los 
indigenas. FE] liquido resultante por la decoccién de las hojas lo toman 
para alucinarse’’. 22-IX-1953, Fernandes 2641 (COL).—Sibundoy. 
““N.v. “Culebra-borrachero’, "Mtzkway borrachera (Kamsa). Arbusto 
hasta de +4 m.; muy folioso. Flores blanecas, péndulas, aromiaticas por 
la noche. Usado pér médicos como narecdético, para emborracharse, 
para resfriados y contra hinchazones. Se reproduce solo por estacas. 
Se dice que rara vez da frutos, que son cilindricos, como un banano,”’ 
12-VI-1956, Idrobo 2221 (COL),—1.5 km. S Sibundoy. ** “Culebra 
borrachera’. Medicinal. Arborescent 3 m.; corolla white; not fruit- 
ing. Indian garden, very infreq.”’ 13-IV-19638, 764 (COL, ECON, 
BISH, Kk, PASTO, S, US).—-Sibundoy. ** ‘Culebra borrachera’, Tree 
3.5 m.:; corolla white, not fragrant, emerging irregularly folded. 
Garden, very infreq.’’ 26-IV-19638, 888 (ECON ).—Sibundoy. ** ‘Cu- 
lebra borrachera’. Tree 3.5 m. ; unopened calyx bulging at top; corolla 
white; filaments often bent, anthers seldom straight; no fruits. Gar- 
den, very infreq.”’ 11-VI-1968, 7772 (COL, ECON, BISH, Kk, 
PASTO, S, US).—-2 km. SW Sibundoy. ** “Culebra borrachera’. Tree 
4 m.; flrs. pendulous, corolla white: corolla and filaments often bent 
irreg. Indian garden, very infreq.’’ 19-IX-19638, 7397 (ECON).— 
1.5 km. S$ Sibundoy. **"Culebra borrachera’. Tree 3 m.; firs. pendu- 
lous, corolla white, bent. Indian garden, infreq.’* 21-IX-1968, 7/400 
(EKCON)—San Francisco. ** “Culebra Borrachera.’ Sparsely branch- 
ing, brittle shrub, 1.5-2.5 m. Flowers white. Infusion of leaves taken 
as hallucinogen. Found only in cultivation.** 22-VIII-1964, Olday 
639 (ECON). 


Datura ‘Andrés’ n. cv. 

Leaves lanceolate to narrowly elliptic, ratio of width 
to length .834-.443; calyx 11.6-18 cm., terminating in 
2-5 lobes; corolla nearly the longest known in the genus, 
31.3-43.2 cm., the slender basal part of the tube exceed- 
ing the calyx, the corolla teeth 8-5.8 em., the limb only 
slightly recurved: anthers distinct, 81-86 mm., stamens 
very long, 25.7-31.6 ¢m., pistil with 1-8 pronounced 


curves in the region of the anthers, exceptionally long, 
32.8-39.8 cm.; not known to fruit. 

Selecting seven characters which most frequently dis- 
tinguish D. candida from D. suaveolens, I find that in 
three, the leaf shape, calyx apex and corolla tooth length, 
‘Andrés’ is intermediate. In the length of its corolla and 
the conspicuous slender basal part of the tube, it is simi- 
lar to D. suaveolens, but the calyx width and free anthers 
are like those of D. candida. The S-curved style is unique 
among the tree Daturas. There seems little alternative 
for the present but to interpret cv. Andrés as a hybrid 
between D. candida and D. suaveolens. 

Common names: Andaqui borrachera. 

Andaqut is the name of a now almost extinct Indian 
tribe to the northeast at the head of the Magdalena Val- 
ley. The area lies many days’ journey away, mostly 
through the eastern lowlands. 

On the assumption that ‘Andrés’ is a hybrid derivative 
of D. suaveolens, a lowland species not known to occur 
in the Valley of Sibundoy, it is probable that it was 
brought to the Valley from the eastern lowlands, or pos- 
sibly directly from the Upper Magdalena area of the 
Andaqui people. In the immediate eastern lowland area, 
Datura is represented by several collections (Klug 1889, 
A, GH, NY, US: Cuatrecasas 10752, US; Schultes 
3472, GH) which resemble ‘Andrés’ but have connivent 
anthers and lack a curved style, and thus approximate 
D. swaveolens more. 

Reference to Datura dolichocarpa (Lagerh.) Safford 
in the Valley of Sibundoy (86) is probably to D. candida 
‘Andrés’. 

Cotombia: Comisaria del Putumayo, Valle de Sibundoy, alt. 2200 
m.—San Andrés. Shrub 2.5 m.; corolla white. Border, infreq. 18- 
IIl-1963, 650 (COL, ECON, PASTO, US).—San Andrés. Arbores- 


cent 3 m.; corolla white; style with S-curve in region of anthers. 


Roadside, infreq. 20-VIII-1963, 1314 (ECON).—2 km. SE Sibun- 
[ 223 ] 


doy. ** ‘Andaqui borrachera’. Tree 2.5 m.; corolla penduluos, white. 
Indian garden, very infreq.’’ 10-X-1963, 7449 (ECON). 


ACKNOWLEDGMENTS 


The investigations upon which this report is based 
were supported by a Public Health Service Research 
Grant (MH 06941-01), by Harvard University and by 
the American Anthropological Association. Much of 
the report was written during the tenure of a Public 
Health Service Postdoctoral Fellowship (1-F2-GM- 
24,938-01). The University of Hawaii provided for the 
illustrations. Aid given by officers of these institutions 
was essential and is sincerely appreciated. 

The following persons have offered suggestions for 
improving the foregoing presentation: W. C. Evans, 
C. B. Heiser, Jr., Linda Howard, R. P. Kahn, N. P. 
Kefford, C. S. K. Raju, R. EK. Schultes and Haydée 
Seyas. I thank each one but do not hold them respon- 
sible. 

I am also grateful to Sharyn Amu for her meticulous 
care in the preparation of drawings for six plates. 


1. 


6. 


“ 


10, 


LITERATURE CITED 


Avery, A.G., 5. Satina and J. Rietsema. 1959. Blakeslee: The 
Genus Datura. Ronald Press, New York. xli+289 pp. 


Barclay, A.S. 1959a. New considerations in an old genus: Datura. 
Bot. Mus. Leafl. Harvard Univ. 18: 245-272. 9 illus. 


. ——. 1959b. Studies in the genus Datura (Solanaceae) I. Tax- 


onomy of subgenus Datura. Doct. dissert., Harvard Univ., Cam- 
bridge. 221 pp. 


Bose, B.C., M.A. Matin, R. Vijayvargiya and M. Lahiry. 1966. 
Effect of solanaceous alkaloids on the 5-hydroxytryptamine con- 
tent of rat brain. Journ. Pharm. Pharmacol. 18: 690. 


Bristol, M.L. 1965. Sibundoy ethnobotany. Doct. dissert., Har- 
vard Univ., Cambridge. 361 pp. 


. 1966a. The psychotropic Banisteriopsis among the Sibundoy 
of Colombia. Bot. Mus. Leafl. Harvard Univ. 21: 113-140. 2 
illus. 


——, 1966b. Notes on the species of tree Daturas. Bot. Mus. 
Leafl. Harvard Univ. 21: 229-248. 4 illus. 


——, 1969. Sibundoy agricultural vegetation. Proc. 37th Intn’l 
Congr. of Americanists, Mar del Plata. (In press. ) 


Castellvi, P. Marcelino de. 1962. Censo indolingiiistico de Co- 


2 


lombia. Amazonia Colomb. Americanista 6, no. 11 extra, 20-34. 


Claus, K.P. and V.E. Tyler, Jr. 1965. Pharmacognosy. Lea & 
Febiger, Philadelphia. 5th ed. 572 pp. 


Evans, W.C. and M. Pe Than. 1962. The alkaloids of the genus 
Datura, section Brugmansia. Part I. D. cornigera Hook. Journ. 
Pharmacy and Pharmacol. 14: 147-156. 


Evans, W.C. and W.J. Griffin. 1963. The alkaloids of the genus 
Datura, section Brugmansia. Part Il. New monotigloyl esters of 
the leaves of D. cornigera Hook. Journ. Chem. Soc, 832: 4348- 
4350. 


Evans, W.C., Valerie A. Major and M. Pe Than. 1965. The 
alkaloids of the genus Datura, section Brugmansia. III. Datura 
sanguinea R. & P. Planta Medica, 13. Jahr.: 353-558. 


[ 225 | 


21. 


we 
co 


Evans, W.C. and Valerie A. Major. 1966. The alkaloids of the 
genus Datura, section Brugmansia. Part IV. New alkaloids of 
D. sanguinea R. & P. Journ. Chem. Soc. (C), 1621-1623. 


. Gottschalk, Werner, 1954. Die Chromosomenstruktur der Solan- 


aceen. Chromosoma 6: 539-626. Illus. 


Heiser, Charles, Jr. 1963. Numeracion chromosémica de plantas 
ecuatorianas. Ciencia y Naturaleza 6, no. 1: 2-6. 


Henry, T.A. 1949. The Plant Alkaloids. Blakiston, Philadel- 
phia. 4th ed. 804 pp. 


Howard, Linda. 1967. Camsa phonology. MS. 
Juajibioy Ch., Alberto. 1962. Breve estudio preliminar del grupo 


aborigen de Sibundoy y su lengua Kamsa en el sur de Colombia. 
Bol. del Inst. de Antropol. (Medellin) 2: 3-38, 9 illus. 


. Juajibioy Ch., Pedro. 1960. Datos recogidos sobre las hojas de 


la planta borrachera. MS., Botanical Museum, Harvard Univ., 
Cambridge. 


Kahn, R.P. and R. Bartels. The Colombian Datura virus—a new 
virus in the potato virus Y group. Phytopathology. (In press. ) 


Kreig, Margaret B. 1964. Green Medicine: The search for plants 
that heal... . Rand McNally, New York. 94-96. 


Lira, J.A. 1945. Diccionario Kkechuwa-Espafiol. Tucuman. 
1200 pp. 


. Ortiz, S.E. 1954. Estudios sobre lingiiistica aborigen de Colom- 


bia. Bogota. 503 pp. 
Pachter, I.J. and Alice F. Hopkinson. 1960. Note on the alka- 


loids of Methysticodendron amesianum. Journ. Amer. Pharm. As- 
soc., Sci. Ed., 49: 621-622. 


Pazos, Arturo. 1966. Glosario de quechuismos colombianos. Im- 
prenta del Departamento, Pasto. 147 pp. 


Perry, Roger. 1963. The Daturas and the Valley of Sibundoy. 
Gardeners Chron. Gardening Illus. Nov. 4. 334-335. 5 illus. 


Safford, W.E. 1921. Synopsis of the genus Datura. Journ. Wash. 
Acad. Sci. 11: 173-189. 3 illus. 


Schultes, R.E. 1955. A new narcotic genus from the Amazon 
slope of the Colombian Andes. Bot. Mus. Leafl. Harvard Univ. 
17: 1-11. 4 illus. 

[ 226 ] 


30. ——. 1961. Native narcotics of the New World. Texas Journ. 
Pharmacy 2: 141-167. 15 illus. 


31. ——. 1963a. Hallucinogenic plants of the New World. Har- 
vard Rev. 1: 18-32. 


32. ——. 19638b. Botanical sources of the New World narcotics. 
Psychedelic Rev. 1: 145-166: in The Psychedelic Reader, Uni- 
versity Books, New Hyde Park, 89-110 (1965). 


33. ——, 1965. Ein Halbes Jahrhundert Ethnobotanik Amerikan- 
ischer Halluzinogene. Zeit. fir Arzneipfl. 13: 125-157. 17 illus. 


34. Seijas, Haydée. 1967. Sibundoy ethnomedicine. MS. 


35. Theilkuhl, J.F. 1957. Introduccion al estudio del Methysticoden- 
dron Amesianum. Dissert., Univ. Nacional, Bogota. 67 pp. 


36. Usecdtegui M., Nestor. 1959. The present distribution of nar- 
cotics and stimulants amongst the Indian tribes of Colombia. Bot. 
Mus. Leafl. Harvard Univ. 18: 273-304. Map; in Rev. Acad. 
Colomb. 11: 215-228. 9 illus. (1961). 


37. van Steenis, C.G.G.J. 1957. Specific and infraspecific delimita- 
tion. Flora Malesiana, ser. 1, vol. 5: clxvii-cexxxiii. 


38. Yepes A., Silvio. 1953. Introduccién a la etnobotanica colombi- 
ana. Publ. Soe. Colomb. Etnol., No. 1: 6-48. 


Qu 


Camsripce, Massacuusetts, JUNE 25, 1969 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


DE PLANTIS TOXICARIIS E MUNDO 
NOVO TROPICALE COMMENTATIONES V 


Virola as an orally administered hallucinogen 
BY 
Ricuarp Evans SCHULTES 


In 1954, the source of a new hallucinogenic snuff from 
the northwest Amazon was identified as the resin of 
several species of the myristicaceous genus V7irola (17). 
The species employed amongst Indians of the Amazon- 
ian region of Colombia were Virola calophylla Warburg, 
V. calophylloidea Markgraf and probably also V’. elongata 
(Benth.) Warburg. 

Additional but often rather vague reports seemed to 
indicate that this same kind of snuff was prepared and 
utilized in the headwaters of the Orinoco in Venezuela 
(15, 27, 29) and to the north of the Rio Negro of Brazil 
(4,8,9, 14,27). Eventually, specimens and photographs 
documented the use of Virola-bark for this purpose in 
that part of Brazil (24). Finally, in 1968, a detailed 
ethnotoxicological study of these myristicaceous snuffs 
(23) showed that the species employed, apparently to 
the exclusion of others, by diverse groups of Waika In- 
dians of northern tributaries of the Rio Negro in Brazil 
is Virola theiodora (Spr. ex Benth.) Warburg. 

In the absence of phytochemical analysis of the snuff 
or of the resin from which it was made, the identity of 
the active principle of Virola calophylla and V. calo- 


[ 229 ] 


phylloidea was not known, and the suggestion that the 
psychoactive properties of the snuff might be attributable 
to myristicine or some component of it was offered (17). 
Recent investigation of Virola theiodora, however, has 
shown that the resin is rich in tryptamines (2). The 
nyakwana snuff of the Waikas, prepared from resin of 
Virola theitodora with no admixture, contains approxi- 
mately 8% of 5-methoxy N, N-dimethyltryptamine, N, 
N-dimethyltryptamine and traces of other tryptamines 
(9), 

Because of similarity of the psychoactivity of the snuff 
prepared from Virola theiodora and that from J’. calo- 
phylla and V’. calophylloidea, there is now every reason 
to presume that the latter two species owe their effects 
to the same or similar tryptamines as those found in the 
former. ‘These compounds have also been indicated in 
an analysis of the bark of V’7rola calophylla (18). 

As a result of recent chemical studies, /’270/a must 
now be added to the list of South American hallucino- 
gens the effects of which are due to tryptamines. These 
tryptaminic narcotics include Anadenanthera peregrina 
and other species of this leguminous genus; J/imosa 
hostilis; Banisteriopsis Rusbyana; Psychotria psychotriae- 
folia; and possibly Justicia pectoralis var. stenophylla. 

Recent field work in the vicinity of Leticia, Colombia, 
on the Amazon River, has uncovered an interesting new 
method of employing myristicaceous resin orally as a 
psychotomimetic agent. 

In February, 1969, I was engaged in collecting for 
phytochemical study several species of Virola along the 
Rio Loretoyacu near Leticia. One of my native assistants 
was a Witoto Indian—Luis Vargas Martinez—who had 
lived in this region for more than 25 years and with whom 
I had worked over a number of years. He is a native of 
El Encanto on the Rio Karaparana, an affluent of the 


[ 230 ] 


PLateE LXII 


DIAGONAL SHADING 
ON INSET INDICATES 
AREA SHOWN 
IN DETAIL BELOW 
. 
< 
ry 


Uutimo 
@RETIRO ENTRE ROS 
} Ti 


a SABANA 
— tA CHORRERA @ Pe R \ 
’ ! 
~ P @ STA, CATALINA ) 
@ CONCEPCION 
aK 
ay . 
we x 
re \ 


) 
y e@EL RET 
a) 4 @PROVIDENCIA 
@FILADELEIA 


Z (wel cen 
‘ SOLEDAD @ 
S 4? 


\ 
8) \ \ \ ENEA Qj @INDOSTAN 
) aa 4 \ y 


—"* A 


—~, os as DIA 


KARAPARANA~IGARAPARANA , | 
REGION Ue ARICA 
of the 

COLOMBIAN AMAZON © wy \ lam 
TEM ng ly fF 
. (ic a4 A VO LGV 


Area of the Colombian Amazon inhabited by the Witoto Indians. 


(231 J 


Rio Putumayo, which, with its parallel river, the Iga- 
raparana, comprises the major centre of the Witoto In- 
dians (5, 12, 20, 25, 26). Asa result of the nefarious and 
merciless exploitations and slaughter of these Indians 
during the wild rubber boom, this region attracted, at 
least fleetingly, world-wide attention in the early part 
of the present century. The past 40 years, with mission- 
ary contacts, have seen a gradual acculturation of the 
Witotos, and many of their beliefs and customs have 
been altered or have disappeared. This acculturation may 
possibly explain our delay in discovering such an inter- 
esting and unique method of preparing and_ utilizing 
Virola-resin as an hallucinogen. 

During the collecting of the bark from several species 
of Virola, this Witoto assistant informed me that the 
Witotos of his father’s generation ate pellets made of 
cumala resin when they wanted to “‘see and converse with 
the little people’*. Cuma/a, a Peruvian name for several 
species of V7zrola, is employed in the Trapecio A mazonico 
—the Leticia area. The assistant informed me that the 
species of Virola which his people employed did not exist 
or was very rare in the Rio Loretoyacu area, but he did 
describe it as a medium-sized tree, slender, growing well 
above the flood-lands on terra alta. The leaf, according 
to the informant, is ‘‘dry’’ to the appearance, measuring 
about six inches in length, with undulating margins. The 
bark, from which copious red resin exudes, is thin, light 
brown. The Witoto name of the tree in the Rio Kara- 
parana is 00-/'00-na. 

While it was impossible for us to collect a voucher 
specimen in the Rio Loretoyacu, I believe that the spe- 
cies represents Virola theiodora. In 1942, amongst the 
Witotos of the Rio Karaparana region, I collected (R. 2. 
Schultes 38878') Virola theiodora with the annotation that 


' Comisaria del Amazonas, Rio Karaparan4, along path from E] En- 


[ 232 ] 


it was ‘‘intoxicating’’. I could get no further informa- 
tion at that time. The Witoto name of this plant in the 
Rio Karaparana was oo-hoo-na, the same name given to 
me by the informant, but this Indian name may refer 
generically to more than one species of Virola or even 
to species in related genera. A collection (22. 7. Martin 
et C.A. Lau-Cam 1272) of Dyalanthera parviflora from 
the Rio Napo in adjacent Peru, for example, bears the 
Witoto name w-Av-na. 

The Witotos rasp the inner part of the freshly stripped 
bark, roll the rasped tissue into balls and express the 
resin from it into a pot of water which is boiled for five 
or six hours, until the mass becomes a thick syrup that 
sticks to the wooden paddle that has been used to stir 
the boiling mixture. 

While this operation is progressing with the resin of 
Virola, another Indian reduces to ashes the bark of a 
‘‘large tree with long leaves, like matamata’’ that grows 
in floodland. We were fortunate in being able to collect 
flowering material of this tree in the Rio Loretoyacu 
region: it represents Gustavia Poeppigiana Berg ex 
Martius.” The Witoto name of this tree is he-rog. 

The ashes are placed in a funnel made of strong leaves, 
and cool water is poured over the ashes and allowed to 
seep through until ‘‘no more cloudiness leaches out’’. 
The water is then slowly boiled down until a greyish 
residue or ‘‘salt’’ is left. This is known in Witoto as 
le-sa. 


canto to La Chorrera. .‘‘Small tree. Height 40 ft. Red resin in bark. 
Intoxicating. Witoto: oo-kod-na’’. May 31-June 2, 1942. R.E. 
Schultes 8878. 

*Comisaria del Amazonas, Rio Loretoyacu, 3 km. above Puerto 
Narifio. In lowland forest. ‘“Tree 7 m. tall. Leaves dark green, 
shiny. Petals white, stamens yellow. Fruit green, seeds germinating 
within ripe fruit.’’ January 28-February 7, 1969. 7. Plowman, T’. 
Lockwood, H. Kennedy et R.E. Schultes 2432. 


[ 233 | 


The thickened JV7rola-resin is then rolled with the 
fingers into tiny pellets the size of coffee beans, and these 
are rubbed in the salt-like residue from the leached-out 
lecythidaceous bark ashes. The pellets, thus coated, are 
ingested whole or dissolved in water and drunk. From 
three to six pellets—called oo-/od-he —are taken initial- 
ly, and the intoxication is said to begin within five 
mintes and last up to two hours. More pellets may be 
taken when the effects of the drug begin to lessen in 
intensity. 

According to my informant’s description, all kinds of 
visual hallucinations are experienced. The narcotic is 
taken usually ina group comprising from three to eight 
or more men, normally including the payé or witch- 
doctor. Only the payé may take the initiative in pre- 
paring the drug. [tis taken not regularly but at irregular 
intervals when the need arises, and only for divination, 
to ‘‘see and converse with the little people’’, to prophecy, 
to find lost property, to ‘‘study’’, to ‘‘talk with’’ people 
from other tribes over great distances and to ensure luck 
in the hunt. 

Dr. Horacio Calle, anthropologist at the Universidad 
Nacional de Colombia, has recently sent me some very 
interesting data and botanical specimens collected from 
informants of the Muinane tribe, now living in the vi- 
cinity of Leticia. These Indians lived originally in the 
Karaparanid-I garaparand-Kahuinari area, the general re- 
gion inhabited by the Witotos. 

According to Calle, the Muinanes call the Viro/a tree 
kutrucu and the drug prepared from its resin Autru. The 
related Bora tribe likewise knows the narcotic which, in 
their language, is hurru. 

Several very young leaves, collected by Calle, are 
difficult to identify as to species. According to Calle, 
botanists at the Instituto de Ciencias Naturales in Bogota 


[ 234 ] 


Pirate LXNIII 


VIROLA 
thetodora 

(Spr: ex Bth,) 

Warb. ug 

| 7] 


Ev 
aps 


tentatively referred them to one of the following species: 
Virola elongata, V. peruviana (A.DC.) Warburg or V’. 
calophylloidea. His Indian informant stated that there 
are several ‘‘kinds of wewna, some with leaves larger than 
others’, but that the tree with the smallest leaves is 

‘the cu ucuna’, 1.e., Meaning probably the Virola 
employed to prepare the hallucinogen. These Virola 
leaves which Calle collected are now preserved in the 
Economic Herbarium of Oakes Ames and appear to me 
possibly to represent /’. calophylloidea.’ 

The data concerning the admixture which Calle re- 
ports are perhaps even more interesting. In preparing 
the pellets for ingestion, the Muinane informant stated 
that the Virola-bark, freshly stripped from the tree, is 
crushed and boiled in water, until the liquid is thick and 
of a honey-like consistency. This syrup is then mixed 
with ashes of the rind of fruit of a species of Theobroma 
or of the dried leaves of a palm, Mawritiella aculeata 
(H Bk.) Burret. The mixture is then rolled into soft, 
brownish pellets. 

It would seem, thus, that there is some variation in the 
methods of preparing the drug. Further field work in the 
original home region of these Indians will be necessary 
fora full understanding of this interesting hallucinogen. 

Interest inthis newly discovered hallucinogen does not 
lie wholly within the bounds of anthropology and ethno- 
botany. It bears very directly on certain pharmacological 
matters, and, when considered with other plants with 
psychotomimetic properties due to tryptamines, this 
new oral drug poses problems which must now be faced 
and, if possible, toxicologically explained. 

It is generally accepted pharmacologically that the 
pure compound N, N-dimethyltryptamine is inactive 


* Comisaria del Amazonas, 8 km. north of Leticia. Muinane name: 
kutruku. April 1969. H. Calle sin. num. 


[ 236 | 


when administered orally, unless accompanied by a mono- 
amine oxidase inhibitor (6). The tryptamines—especially 
N, N-dimethyltry ptamine—are the active constituents 
of hallucinogenic snuffs of South America (2, 6, 18, 22, 
27) prepared from Anadenanthera peregrina (yopo), 
A. colubrina (huilea) and Virola thetodora and probably 
other species (yakee, epend, nyakwana, paric’) (22). Yet 
the narcotie vinho de yurema, prepared from the roots of 
Mimosa hostilis (11), the active constituent of which is 
N, N-dimethyltryptamine, is extremely active when 
taken orally. It has furthermore recently been learned 
that the two common admixtures of the ayahuasca, caapi 
or yajé drink prepared basically from bark of either Banis- 
teriopsis Caapi or B. inebrians, rich in harmala alkaloids, 
are leaves containing N, N-dimethyltryptamine: Banvs- 
teriopsis Rusbyana (1, 6, 16) and Psychotria psychotriae- 
folia (7). There is no question but that the addition of 
leaves of Psychotria psychotriaefolia and especially of 
Banisteriopsis Rusbyana very strongly enhances the color 
visions, increases the intoxication and prolongs the nar- 
cotic effects of the drink. In the case of these two ad- 
mixtures, it is clear that, when added to a beverage 
containing the harmala alkaloids, harmine and harmaline, 
they would be in the presence of monoamine oxidase 
inhibitors. As has recently been suggested, ‘*. . . perhaps 
this fact, coupled with the presence of small amounts of 
related isomers, may help account for the effectiveness 
of the crude preparation. . .”’ or ‘‘perhaps the strong 
concentration of active principles or the presence of other 
unidentified substances facilitate absorption”’ (6). 

In the instance of yurema, which is prepared appar- 
ently from the roots of Mimosa hostilis with no admix- 
ture whatsoever, a monoamine oxidase inhibitor might 
possibly be present, but we do not yet know. With the 
orally administered Virola-resin of the Witotos, one 


[ 287 | 


must assume that the tryptamine acts without such an 
inhibitor or else that the residue from the leaching of the 
lecythidaceous bark-ashes contains a monoamine oxidase 
inhibitor. 

All of this points out with singularly insistent empha- 
sis the wisdom of an interdisciplinary approach in the 
search for new psychoactive drugs, since the value of 
folklore data has so often in recent studies been indicated 
(18). It points further to the urgency of intensive ethno- 
toxicological study of folklore in view of the rapid dis- 
integration and disappearance of aboriginal cultures 
around the world (19, 21). 

The field work during which the discovery reported in 
this paper was made was financially supported in part by 
the Bache Fund of the National Academy of Science, 
to which source grateful acknowledgment is herewith 
offered. 

This paper was read at the 10th Annual Meeting of 
the Society for Economic Botany, Longwood Gardens, 
Kennett Square, Pennsylvania, April 14, 1969. 


[ 288 | 


10. 


11. 


13. 


14, 


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148-151. 


. Agurell, S., B. Holmstedt, J.-E. Lindgren and R.E. Schultes. 


** Alkaloids in certain species of Virola and other South American 
plants of ethnopharmacologic interest’? Acta Chem. Scand., in 


press. 


Altschul, S. von R. ‘‘Vilea and its use’? in ‘‘Ethnopharmacolo- 
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Becher, H. ‘‘Die Suraraé und Pakidai, zwei Yanonami Stamme 
in nordwestbrasilien’’ Mitteil. Mus. Vélkerkunde Hamburg 26 
(1960) 1-138. 


Collier, R. “‘The river that God forgot’’ (1968). 


Der Marderosian, A., H.V. Pinkley and M.F. Dobbins. **Native 
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Banisteriopsis Rusbyana’’ Am. Journ. Pharm. 140 (1968) 1387-147. 


Der Marderosian, A. Personal communication. 
Ducke, A. Arch. Inst. Biol. Veg. 4 (1937) 3. 
—., **As leguminosas da Amazonia brasileira’’ (1939) 41. 


Farnsworth, N. and K. Euler. “‘An alkaloid screening procedure 
utilizing thin-layer chromatography’’ Lloydia 25 (1962) 186-195. 


Goncalves de Lima, O. ““Observacoes sobre o ‘vinho de jurema’ 
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Inst. Pesqu. Agron. 4 (1946) 45-80. 


Hardenberg, W.E. ““The Putumayo—the devil’s paradise”’ 
(1912). 


Holmstedt, B. and J.-E. Lindgren. ““Chemical constituents and 
pharmacology of South American snuffs’’ in ““Ethnopharmacolo- 
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Knobloch, F. “‘Die Aharabu-Indianer in Nordwest-Brasilien”’ 
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18. 


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(a3) 
to 


23. 


28. 


29. 


Koch-Grinberg, T. **Zwei Jahre unter den Indianern’’ 1 (1909) 
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—., ‘‘The role of the ethnobotanist in the search for new 
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‘ . oe : : 6 yy 
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and B. Holmstedt. ‘‘De plantis toxicariis e Mundo Novo 
tropicale commentationes II. The vegetal ingredients of the 
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Seitz, G.L. ““Epena, the intoxicating snuff powder of the Waika 
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Casement’’ (1959) 208-315. 


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American Indians’’ 3 (1948) 749-762 [Bur. Am. Ethnol. Bull. 
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Wassén, S.H. and B. Holmstedt. ““The use of parica, an ethno- 
logical and pharmacological review’’ Ethnos 28 (1963) 5-45. 


Whiffen, T.W. ““The northwest Amazons”’ (1915). 


Wilbert, J. “‘Indios de la regién Orinoco-Ventuari’’ Fund. La 
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[ 240 ] 


BOTANICAL MUSEUM LEAFLETS 


HARVARD UNIVERSITY 


CampripGcr, Massacnuserrs, NoveEMBER 21, 1969 VoL, 22, No. 


A NEW AMAZONIAN ARROW POISON: 
OCOTEA VENENOSA 
BY 
A.J. KosTeERMANS,’ Homer V. PINKLEY,* 
AND WiLu1AM L. STERN ’® 


A RECENTLY discovered ingredient of arrow poisons used 
by the Kofin Indians of the westernmost Amazon in 
Colombia and Ecuador is produced by an undescribed 
species of Ocotea of the Lauraceae. 


Ocotea venenosa Kosterm. et Pinkley sp. nov. 

Arbor ramulis cum innovationibus dense minutissime 
ferrugineo-pilosis; foliis spiraliter alternantibus vel sub- 
oppositis, chartaceis, ellipticis, obscure acuminatis, basi in 
petiolum brevem contractis supra laevibus, nervo medi- 
ano prominulo, costis filiformibus vix prominulis, subtus 
dense prominule reticulatis, nervo mediano prominent, 
dense pulverulente piloso, costis utrinque ca. 9 subpaten- 
tibus prominulis; inflorescentiis paniculatis, axillaribus, 
parvis, dense minutissimeque ferrugineo-pilosis; floribus 
tantum in alabastrum cognitis; tepalis ovatis ; staminibus 
fertilibus 9, crassis, exterioribus loculis 4 magnis, in- 
trorsis, filamentis crassis, interioribus loculis 4 extrorsis ; 
glandulis non vidi; ovario subpiloso, ellipsoideo, in sty- 


' Herbarium Bogoriense, Bogor, Indonesia. 
? Botanical Museum of Harvard University, Cambridge, Mass. 
* Department of Botany, University of Maryland, College Park, Md. 


(241 J 


lum brevem prodeuntibus; fructu depresso-globoso, 
magno in cupulo magno incluso. 

Cotompra: Comisaria del Putumayo, Rio Guamites, Santa Rosa. 
Primary forest. Altitude about 1060 feet. Long. 77°05’ W, Lat. 
00°19’ N. November 26, 1966. H.V’. Pinkley 555 TYPE! (Type in 
Herb. Gray. Dupticare typrk in Econ. Herb. Oakes Ames; Utrecht; 
Naturhistoriska Riksmuseet, Stockholm).—Ecuapor: Provincia del 
Napo, Rio Aguarico, Durero. ““Fruit used to make arrow poison. 
Wood, yellow, very hard and bitter to taste. Sterile’’. October 26, 
1966. H.V. Pinkley 538, 

Tree 32 m., bole 20 m., diam. 70 em.: branchlets and 
terminal bud densely and minutely rusty-pilose. Leaves 
spirally arranged, occasionally subopposite, chartaceous, 
coriaceous, glabrous except above the midrib underneath, 
elliptical or narrowly elliptical, 6-9 16-20 cm. obscure- 
ly acuminate, base contracted into petiole 8-12 mm. 
long, flattened above; upper surface smooth, glossy, 
midrib conspicuous, prominent, lateral nerves filiform, 
hardly prominent; lower surface glossy, densely, mi- 
nutely, and rather obscurely reticulate. Midrib promi- 
nent, densely pulverulent-pilose, lateral nerves ca. 9 
pairs, rather patent, slightly arcuate, prominent. Pani- 
cles (immature) axillary, up to 2 cm. long, densely, 
minutely rusty-pilose. Flower buds ca. 1.5 mm. in diam. : 
tepals ovate, fleshy, acutish, equal, or the inner ones nar- 
rower. [Fertile stamens 9, thick, 4-celled; outer ones 
with large introrse and introrse-lateral cells, the anther 
not differentiated from the thick filament; the inner row 
with similar but narrower stamens with extrorse cells: 
the connectives blunt, protruded beyond the cells. 
Glands not seen. Ovary and style as long as the stamens, 
the ovary ellipsoid, merging into a slightly shorter style 
with rather inconspicuous stigma. Fruit depressed- 
globose, 5.5 cm. diam., 4.5 em. high, smooth, with a 
small notch at the apex; cup 1 mm. thick, obscurely, 
broadly ribbed, covering the entire fruit except the up- 


[ 242 ] 


per part, leaving an orifice of 2.5 cm. diam.; the cup 
closely adpressed to the fruit. In a smaller fruit (4.5 em. 
diam.) the orifice of the cup is 3.5 cm. in diam. Stalk 
thick, woody, very short, 6 mm. diam.; cotyledons 
plane-convex, thick; plumule, and radicle basal. 

Most of the flower buds available for dissection were 
abnormal as a result of fungal attack. The fruit cup is 
unusual in Ocotea, because it is almost adnate to the 
fruit. Another peculiarity is that the fruit does not pro- 
trude from the cup. This is rare in Ocotea, but it does 
occur in other genera of the Lauraceae. 

The Kofiin Indians, a tropical rain forest tribe of east- 
ern Ecuador and Colombia, employ the fruit of this plant, 
gingivé’k'o in Wofain, as an ingredient in one of their 
arrow poisons. Hence, the specific epithet, which means 
‘*very poisonous’’ in Latin, was chosen for this large tree. 
Two trees had to be felled before fertile material was 
found. After felling the first tree, which was sterile, I 
(Pinkley) took bark of the tree to my hut, thinking that 
it was the part of the tree used in preparing the arrow 
poison. When L arrived with the bark, the Indians laughed 
and revealed that they use only the fruit. 

Chemical analysis and pharmacologic research of this 
new species is being carried out by Dr. Ara Der Mar- 
derosian of the Philadelphia College of Pharmacy and 
Science. He reports, through personal communication, 
that he has isolated two of several alkaloids present, 
rodiasine and demethylrodiasine related to the curare 
alkaloid of medicine, d-tubocurarine. Rodiasine was first 
isolated in crystalline form from Ocotea Rodiei (Rob. 
Schomb. ) Mez (Nectandra Rodiei Rob. Schomb.), (4, 7). 
The alkaloid study of Ocotea Rodiet began much earlier, 
however, when Maclagan in 1848 separated the alkaloids 
of this plant into two amorphous fractions, calling the 
portion soluble in ether bebeerine, and the portion in- 


[ 243 | 


soluble in ether sepeerine. Since Maclagan’s classic study, 
the name bebeerine, however, has been variously defined 
resulting in much confusion (7). Since he isolated amor- 
phous fractions rather than crystalline compounds, most 
are in favor of abandoning his nomenclature and using 
coined names referring only to actual isolated compounds. 
Hence, though Maclagan’s bebeerine may be, in fact, the 
same or related to rodiasine, the term rodiasine is now 
preferred. Maclagan was encouraged to study the chem- 
istry of Ocotea Rodie: not because of any known curariz- 
ing effect but because the wood was known to be highly 
resistant to insects, marine borers and fungal decay (5,8). 
Today we know that curine, the / form of ‘‘bebeerine,”’ 
in high concentration ‘‘. . . causes paralysis of the stri- 
ated muscles and paralysis of the nerve end plates”’ (6). 
Though the chemical constituents of Ocotea venenosa and 
Ocotea Rodie: appear to be similar, the initial chemical 
investigations were induced for two different reasons. 
Several plants of the Lauraceae with an alkaloid related 
to ‘“‘bebeerine’’ were reported in 1890 by Greshoff, a 
pioneer in phytochemistry. Greshoff found in several 
genera an alkaloid which he called ‘‘lauro-tetanine.’” He 
stated that (transl.), ‘‘according to the important charac- 
teristic, . . . causing tetanus in several species of animals, 
I propose to give this body which can be crystallized and 
characterized by good reactions the name lauro-tetanine’’ 
(3). He furthermore recognized the similarity of lauro- 
tetanine and bebeerine and suggested that (transl.) ‘‘a 
detailed quantitative research of bebeerine seems one of 
the most urgent desiderata of the alkaloid studies’’ (8). 
Other alkaloids with curarizing effects have been iso- 
lated from the lauraceous tree Cryptocarya Bowier 
(Hooker) Druce of Australia (2). The poisonous nature 
of this plant was discovered, quite by accident, by 'T. L. 
Bancroft in 1886. He related his experience: (1) 


[ 244 | 


Puate LXV 


OCOTEA venenosa- Kosterm. ef 


Pink/ley 


. 
mans == 


nin C19 MT OS 


Ocotea venenosa. 
Qo 


Habit sketch of branchlet, 2 
2a, outer stamen, 


1, inflorescence branchlet, 
Por 


15, 2b, inner stamen, 3, ovary and style, 


20. 
t, diagrammatic cross section of fruit, 3. 5, diagrammatic longitudinal section 
of fruit, ». 6, oblique angle of dissected fruit showing large cotyledon and 
developed root, » }. 7, fruit, ~ 4. 


On May 12th, 1886, in search of poisonous plants | found the 
bark of this tree to have a very persistently bitter taste. Physi- 
ological experiments were immediately made, which led to the 
discovery of its toxic action. Other species of the same genus are 
likewise poisonous. It is interesting botanically to note such a 
poisonous genus in this order. 

The alkaloid or its salts have an intensely bitter taste; it is 
odourless and extremely poisonous, slightly soluble in water, 
very soluble in alcohol, ether, and chlorotorm. Warm blooded 
animals poisoned with Cryptocarya exhibit respiratory difhculty, 
soon ending in asphyxial convulsions and death. On frogs it 
causes paralysis of the reflex function of the spinal cord and the 
peripheral ends of motor nerves as effectually as curara. 


The fact that Bancroft was led to make further studies 
because of the very persistent bitter taste has an interest- 
ing parallel among the Kofin Indians. Most, if not all, 
of the plants which they use in preparing arrow poisons 
are bitter. Mqually true, one of the first diagnostic charac- 
ters used by the Nofins in identifying a plant is its taste. 
After first tasting the bark of a tree, for example, they 
will look up into the forest canopy and try to find leaves 
associated with the tree. It is quite possible that bitter- 
ness, inza/tst in Kofan, led to the original discovery of 
the plants used for their arrow poisons. Moreover, in 
many cultures bitterness has become associated with 
death. An association which may have resulted from 
primitive man’s observation that many bitter plants can 
‘cause death. 


NYLEM ANATOMY OF Ocotea venenosa 


Pores are decidedly rounded and occur mostly in the 
solitary (76% ), radial multiple (20% ), and clustered (4%) 
distributions. Radial multiples usually comprise only 2 
pores, 8- and 4-pored multiples being rather uncommon. 
Only 8-pored clusters were noted. Tangential pore 
diameter ranged from 87—-155p with an average of 117p, 
based on measurements of 50 pores. Vessel elements all 


[ 246 ] 


1, radial section of xylem showing fenestriform pitting between vessel elements and xylem parenchyma, 290. 


2, coronated vessel element from macerated material illustrating fenestriform pits at ends, “140. Note: Perfora- 
tions in this vessel element are out-of-focus. 


ALV Iq 


AX'T 


contain simple perforations. Occasional vessel elements 
with three perforations were noted. Ligules may be 
present and are usually short or they may be absent from 
the ends of vessel elements. Intervascular pitting is al- 
ternate; inner apertures of pits are horizontally elongate 
and are enclosed by borders which may be elongate, 
rounded, or polygonal; adjacent inner apertures may be 
crossed or coincident. Vessel elements range in length 
from 2354—-9385p and average 564m in length, based on 
50 measurements. 

The relationship between vessel elements and axial 
and ray parenchyma cells presents a curious if not unique 
situation in this species. Both kinds of cells are intercon- 
nected with adjacent vessel elements through unilaterally 
compound pitting with very large simple pits in paren- 
chyma cells subtending two or more bordered pits in 
vessel elements. The large simple pits in the parenchyma 
cells are often traversed by branched or simple filiform 
processes of cell wall material. In addition to the uni- 
laterally compound pitting, half-bordered pits occur. 
Supplementing these more or less common kinds of pit- 
ting is the pitting in the coronated vessel elements. 
These vessel elements are normal in appearance in all 
respects except that upper and lower ends bear a com- 
plete or partial ring of large, fenestriform, obscurely 
bordered pits which give the impression of a crown or 
corona (Plate LXIII, fig. 2). These are associated with 
simple pits of similar size and form in adjacent ray and 
axial parenchyma cells (Plate LXIII, fig. 1). Pits of the 
corona may be square, rectangular, triangular, oval, el- 
liptical, or irregular. At times these pits occur on ligules 
as well. The uniqueness of this pitting resides in its posi- 
tion at upper and lower ends of vessel elements, its vessel 
element encircling tendency, and the large size of the 
pits. All combine to impart a crown-like appearance to 
the ends of vessel elements 

[ 249 ] 


For all intents and purposes, the imperforate tracheary 
elements are libriform wood fibers. Inner apertures of 
pit-pairs are somewhat elongated and outer apertures are 
nearly circular. A minute, insignificant border may be 
observed under high magnification in some pits of some 
fibers. Walls are very thick ranging up to 8u. Lumina 
are equal to or less than the diameter of the walls and 
are often completely occluded by the growth of the 
secondary cell walls. Fiber length ranges from 750p- 
1437 and averages 1104, based on 50 measurements. 

Vascular rays range from 1—4 cells wide, 1-seriate and 
4-seriate rays being rare. Ray height ranges from one 
cell to over 25 cells, most rays being lower than 15 cells 
high. Rays are heterocellular with the terminal ray cells 
being square, squarish, or upright and with the body of 
the ray comprising only procumbent cells. Terminal ray 
extensions are ordinarily one or two cells high with uni- 
seriate extensions rarely reaching five or six cells high. 
These cells are slightly swollen as viewed in tangential 
section. 

Axial parenchyma is always paratracheal and may con- 
sist of a vasicentric sheath one to several cells wide around 
vessels and vessel groups; it may be aliform with short, 
broad wings; or sometimes it is even aliform-confluent. 

No secretory cells were observed in the secondary 
xylem. 

Because of the unusual nature of some of the morpho- 
logical features of Ocotea venenosa, namely, the fruit, the 
third author felt that an investigation of the secondary 
xylem might be instructive. Other than the exceptional 
coronated vessel elements described above and the some- 
what unusual absence of secretory cells, at least in the 
specimen examined, the xylem of O. venenosa falls well 
within the range of the xylem anatomical characteristics 
of Lauraceae described by Stern in 1954 (9). In that 


[ 250 | 


work, several species of Lauraceae were noted as lacking 
secretory cells in the secondary xylem, namely : Dehaasia 
triandra, Lindera Benzoin, Nectandra coriacea, N. glo- 
bosa, Neolitsea Levinei, and Ravensara crassifolia. Secre- 
tory cells were described as scarce in Ocotea palmana, 
Hypodaphnis Zenkeri, and Laurus nobilis. The absence 
of secretory cells in the secondary xylem of O. venenosa 
cannot alone be used to rule out or render questionable 
the demonstrated taxonomic affinities of this species. 

The peculiar fenestriform pitting in vessel elements 
represents a unique specialization occurring only in this 
particular species, at least as far as is presently known. 
Whether or not the combination of the unusual fruit 
(for Ocotea, anyway) and the special coronated vessel 
elements are significant enough characteristics upon 
which to base a new genus of Lauraceae, will depend 
upon the judgment of taxonomists. 


[ 251 ] 


BIBLIOGRAPHY 


Bancroft, T.L. 1887. On the Physiological Action of Cryptocarya 
australis. The Proceeding of the Royal Society of Queensland. 
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Ewing, Jean, et al. 1953. The Alakloids of Cryptocarya Bowiei 
(Hook.) Druce. Australian Journal of Chemistry. Vol. 6, pp. 78- 
85. 


Greshoff, M. 1890. Eerste Verslag van het Onderzoek Naar De 
Plantedstoffen van Nederlandsch-Indie. Mededeelingen uit 
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Grundon, M.F., and J.E.B. McGarvey. 1960. Alkaloids from 
Greenheart. Part I. The Isolation of the Alkaloids, and the Struc- 
ture of Sepeerine. Jour. Chem. Soc., pp. 2739-2745. 


Maclagan, Douglas. 1843. Ueber den Bebeerubaum des brittischen 
Guiana. Annalen der Chemie und Pharmacie. Vol. XLVIII, pp. 
106-121. 


Manske, R.H.F. and H.L. Holmes (eds.). 1954. The Alkaloids, 
Chemistry and Physiology, Vol. IV. Academic Press, Inc. N.Y., 
P.- 231: 


McKennis, Herbert Jr., etal. 1956. Isolation of a Tetramethoxy- 
lated Alkaloid from Demerara Greenheart. Jour. Amer. Chem. 
Soc. Vol. 78, pp. 245-248. 


Record, S.J. and R.W. Hess. 19438. Timbers of the New World. 
Yale Univ. Press, New Haven, pp. 211-213. 


Stern, W.L. 1954. Comparative Anatomy of Xylem and Phylo- 
geny of Lauraceae. Tropical Woods. Vol. 100, pp. 1-73. 


[ 252 ] 


BLANCHE AMES AMES 
(1878-1969) 


AN APPRECIATION 


TurouGu most of her long life, Blanche Ames Ames 
cherished a unique and meaningful relationship with the 
Botanical Museum, a closeness severed only by death at 
her home in Borderland, North Easton, Massachusetts, 
on March 1, 1969. 

Wite and lifelong helpmeet of Professor Oakes Ames, 
second director of the Botanical Museum, Blanche Ames 
never held an official appointment as a member of the 
Museum staff. Nevertheless, her contributions to the life 
of the institution were so profound that they will always 
be appreciated in the educational and research activities 
of the Museum and, through it, will continue to be 
widely recognized in the world of botany. 

Mrs. Ames was an early champion of women’s suf- 
frage, a leader in civic and charitable affairs and an author ; 
but perhaps she will be most widely acknowledged as an 
artist. Her botanical etchings and pen and ink drawings 
are displayed in the Botanical Museum and in the Metro- 
politan Museum of Art, and her oil paintings hang at 
Harvard, Dartmouth, Columbia, Phillips Exeter Acad- 
emy, the Mississippi State Hall of Governors and in 
several private collections. 

Her interest in botanical artistry began when, at the 
Ames Botanical Laboratory at North Easton, she started 


[i ae] 


to illustrate her husband's publications on new species of 
Philippine orchids. When Professor Ames transferred his 
activities to the Museum and installed there the Ames 
Orchid Herbarium and Library, she intensified her ef- 
forts, producing over a period of some fifty years literally 
hundreds of outstanding line drawings of orchids. When 
the University added the supervision of the Arnold 
Arboretum to Professor Ames’ duties, Mrs. Ames un- 
dertook the preparation of drawings of a number of 
horticulturally important shrubs. 

In the 1920°s, Professor Ames turned his attention 
to economic botany, and his wife began to illustrate 
plants of use to man. Especially noteworthy was Mrs. 
Ames) preparation, under her husband’s guidance, of 
the now famous **Ames Charts of Economic Plants’’. 
The teaching of Harvard University’s course in Kconom- 
ic Botany, offered for many years by Professor Ames, 
was organized around these four colorful wall charts de- 
picting Important economic species arranged on a ‘*phylo- 
genetic tree’, in accordance with the Engler-Prantl 
System. Still in use to-day, the charts have helped in- 
terpret: plant classification to hundreds of students for 
more than fifty years and have stamped a unique charac- 
ter on Harvard's teaching in this interdisciplinary field. 

When Protessor Ames established a press in- the 
Museum in the early 1980’s and initiated publication of 
the Botanical Museum Leaflets, Harvard University 
and occasional books, Mrs. Ames’ artistry vitalized the 
research papers that were printed. Among the books 
produced on this press and illustrated by Mrs. Ames are 
“Orchids in’ Retrospect’*, ‘‘Drawings of Florida Or- 
chids”* and **Economie Annuals and Human Cultures”’. 
Her line drawings have likewise been widely reproduced 
in many monographs, floras and manuals issued by other 
botanical institutions and by commercial publishers. 


[ 254 | 


Mrs. Ames was an accomplished portrait artist. Ex- 
amples of this aspect of her artistic contribution, hanging 
in the Botanical Museum, consist of life-sized oil paint- 
ings of the Museum’s three directors: Professor George 
Lincoln Goodale, Professor Oakes Ames, and Professor 
Paul C. Mangelsdort. [t is unique to have the portraits 
of three men whose service spanned a period of 79 years— 
from 1888 to 1967——painted by a single artist. 

Truly a great lady and an outstanding artist, Blanche 

Ames? influence will long be felt in botany, for she spent 
au great part of her life interpreting the beauty of plants 
for others. This characteristic of her life was appropriately 
stressed at memorial services for Mrs. Ames by the Rev. 
Mr. Edmund Palmer Clarke of the Unitarian Church of 
North Easton. “If Mrs. Ames had been a man, we 
would have said of her that she was ‘a man of parts’... 
wv person of much ability and many talents... Those of 
you who knew her best will see the aptness of my using 
the phrase to describe her. Perhaps even better, now that 
she has gone from us, is Shelley’s...: ‘She is a portion 
of the loveliness which once she made more lovely. 
For this was her greatest talent —to know nature—to 
reproduce it in her drawings for those less perceptive than 
she, and to give expression to those things which made 
us better because she had lived amongst us.”” 

The staff of the Museum can think of no better way 
of expressing its appreciation of Blanche Ames* devotion 
to botanical art than to present, on the very press that 
has known so well her skillful pen, a selection of her 
artistic and scientific contributions. 


—Ricnarp Evans SCHULTES 


PLATE LXVI 


BLANCHE AMES AMES 


yp | 
2I0 | 


PLuatrE LNVII 
ii) 


ih 


‘\ id 
i> 


Te i 
(i 


Davidia involucrata Ball, 
(Dove Tree) 


[f ocr | 
ro | 


Ll 


LNVITI 


PLATE 


Ih 


rardy 


indent (Lindl) € 


sl TC 


I 


adicion 


Polyt 


LNAI. 


. 
4 


PLATE 


ir 


2 
v. 


) 


ish, 


So l / 


s ( 


hellu 


on pulc 


ulopog 


. 
c 


C 


LXNX 


PLATE 


Habenaria nivea (Natt.) Spreng. 


PLateE LNXI 


stig 


] 
i 
aI 
al 
=| 


Cypripedium Calceolus LZ. 
var. pubescens (IW72/d.) Correll 


PiarE LXANII 


pidendrum tampense Land. 


Pirate LNNITI 


Blanche Ames Ames and Oakes Ames 


| 263 


PLATE LXXIV 


avi 


of teh pendulispica Ch mes 


FOUR NEW SPECIES OF SAURAUIA FROM 
SOUTH AMERICA 
BY 
Duasa D. Sorsarto | 


Saurauia is a widespread genus of the Actinidiaceae 
(Theales). The members of Sawrauia are represented in 
both tropical and subtropical Asia and America. Ac- 
cording to the latest estimate made by Melchior,* ap- 
proximately 280 species of the genus have been described. 
The American distribution of Saurauia extends from 
central Mexico in the north to Bolivia in the south, 
through Andean South America. Twenty-two species 
are found in Mexico and Central America,’ and my 
present study * indicates that 49 species are represented 
in South America. Members of the genus consist mostly 
of localized populations of mountainous and subalpine 
trees and shrubs, but species adapted to lower elevations 
and species with broad geographic distribution are also 
known. 

‘Latin American Teaching Fellow, Associate Professor, Departa- 
mento de Biologia, Facultad de Ciencias y Humanidades, Universidad 
de Antioquia, Medellin, Colombia. 

” In Engler, Syllabus der Pflanzenfamilien, ed. 12, 2: 160. 1964. 

* Hunter, G.E. 1966. Revision of Mexican and Central American 
Saurauia (Dilleniaceae). Ann. Missouri Bot. Gard., 53 (1): 47-89. 

* Studies of South American Saurauia (Actinidiaceae), 458 pp., 61 


pls., Ph.D. dissertation (1968), Department of Biology, Harvard 
University, Cambridge, Mass., U.S.A. 


[ 265 ] 


Among the collections of South American Saurauia 
kindly loaned to me for study by the United States 
National Museum, the Gray Herbarium of Harvard 
University and the New York Botanical Garden, there 
were specimens which I could not identify with any of 
the species that [ accepted from South America nor with 
any of those of Mexico and Central America accepted 
by Hunter. After careful studies of these specimens, I am 
inclined to believe that these species are new to science. 


1. Saurauia multinervis Soejarto sp. nov. 


Species foliis magnis elongatis glabris, nervis laterali- 
bus numerosis, inflorescentiis omnino glabris multifloris, 
sepalis glabris notabilibus.—Ho.orypus: Idrobo & Fer- 
nindez 54 (US). 

Shrubs to 2 m., many-branched, glabrous. Leaf blades 
elongate-obovate, cuspidate at apex, broadly cuneate to 
obtuse at base, dentate-serrate with minute and sharp- 
pointed serrations along margins, 80-42 cm. long, 12-14 
ecm. wide, coriaceous, dirty dark olive-brown above in 
dry state, brownish olive beneath, laevigate and smooth 
above, secondary veins 35-87 pairs, tertiary veins ele- 
vated, more prominent than lesser venation, both surfaces 
glabrous; petioles 1-2 ecm. long, 3-5 mm. in diameter, 
glabrous. Inflorescences straight, ca. 150-flowered, 23 
cm. long, 15 em. wide, glabrous, primary peduncle 1.5 
em. long, bracts linear, to 5 mm. long. Flowers un- 
known. Fruits laxly distributed, berries 5-locular, glo- 
bose, to 6 mm. in diameter, 5-sulcate, glabrous; pedicels 
4-8 mm. long, bracteoles linear, to 3 mm. long; persis- 
tent sepals 5, oblong-elliptic to ovate, obtuse, 5-6.5 mm. 
long, 8-4.5 mm. wide, all glabrous within and without, 
marginally entire to irregularly ciliolate ; persistent styles 
5, 4.5-5 mm. long, stigmas capitate. 


[ 266 ] 


Distribution: Colombia (Departamento Cauca), alt. 
2000 m., fruiting in August. 


Vernacular names: Lulumoco (Cauca—Idrobo & Fer- 
nindez). 


Specimens examined. Colombia, Cauca: Cordillera 
Occidental, vertiente oriental, EK] Tambo, Corregimiento 
de Chapa, Ciruelas, Idrobo & Fernandez 54 (US). 


». Saurauia Schultesiana’ Soejarto sp. nov. 

Species ceteribus differt foliis et inflorescentiis ramuli 
apice confertis, laminis supra distante et sparse setulosis 
subtus sparse setosis, inflorescentiis paucifloris (minus 
quam vigintifloribus), bracteis glabris, pedicellis longis 
(usque ad 10 mm.), floribus magnis (20-25 mm. latis), 
sepalis parte mediana parse pubescentibus sed lateralibus 
glabris, intus omnino glabris, staminibus viginti-quinque 
ad trigintiquinque.—Holotypus: Pennell 10501 (NY). 

Trees: copiously to sparingly pubescent. Branchlets te- 
rete, abundantly pulverulent, rusty strigose to appressed- 
setose (young leaves and shoot deep brown). Leaves 
crowded near tip of branchlets; blades elliptic to oblong- 
obovate, acuminate at apex with acumen to 15 mm. long, 
cuneate at base, serrulate along margins, 14-18 cm. long, 
7-8 cm. wide, subcoriaceous, in dry state dark dirty 
brown above, greyish olive-brown beneath, scarcely 
scabrous above and beneath, secondary veins 15-18 pairs, 
tertiary veins elevated, more prominent than lesser vena- 
tion, sparingly setulose along and between minor veins 
and abundantly appressed-setose to strigose along major 
veins above, sparingly setose to setulose (mixed with 


°T have named this species in honor of Dr. Richard Evans Schultes, 
Executive Director of the Botanical Museum of Harvard University 
and my former academic adviser, who has inspired in me a deep in- 
terest in the tropical American flora. 


[ 267 ] 


scattered radiate trichomes) along and between minor 
veins and abundantly pulverulent strigose along major 
veins beneath; petioles ca. 2 cm. long, 2-3 mm. in 
diameter, half-terete, abundantly rusty strigose. Inflo- 
rescences distributed near tip of branchlets, straight, 
6-20-flowered, 13-30 cm. long, 2-7 cm. wide, abundantly 
pulverulent scurvy-strigose, primary peduncle 8-15 cm. 
long, bracts broadly triangular to suborbicular, 10-15 
mm. long, 5-7 mm. wide, glabrous. Flowers 20-25 mm. 
broad, buds to 9 mm. in diameter, pedicels 5-10 mm. 
long, bracteoles broadly triangular, to 10 mm. long: 
sepals 5, outer 2 trullate to oblong-ovate, subacute, 10— 
12 mm. long, 6-7 mm. wide, imbricate one suborbicu- 
lar, rounded, 10-12 mm. long, 8-9 mm. wide, inner 2 
suborbicular to orbicular-oblong, rounded, 10-14 mm. 
wide, all glabrous to medially sparingly strigillose with- 
out, completely glabrous within, marginally entire to 
apically ciliolate; petals 5, white, subquadrangular to 
spatulate, rounded, 15-19 mm. long, 9-11 mm. wide; 
stamens 25-85, filament 4-5 mm. long, anther 2.5-3 
mm. long; ovary 5-7-locular, globose, 5—7-sulcate, gla- 
brous, styles 5-7, to 7.5 mm. long, stigmas capitate. 
Berries 5-7-locular, globose, to 6 mm. in diameter (im- 
mature) 5—7-sulcate. 


Distribution: Colombia (Departmento Caldas), mossy 
forest, alt. 3200-3400 m., flowering in September. 


Specimens examined. Colombia, Caldas: Cordillera 
Occidental, Cerro Tatama, Pennell 10501 (NY). 


3. Saurauia Mexiae® Killip ex Soejarto sp. nov. 
S. pseudostrigillosae Buse. affinis sed floribus grandiori- 
*Killip annotated the collection (Mezxia 8488) as Saurauia Meziae 


Killip spec. noy., but so far as I am aware, the name and description 
of this concept have never been published. 


' 268 J 


bus, pedicellis longioribus, foliis subtus abundanter et 
molliter stellatis differt. —Holotypus: Meawia 8488 (US). 

Trees to 10 m.; copiously pubescent. Branchlets 
somewhat terete, distinctly scarred, abundantly to spar- 
ingly pulverulent strigose. Leaves crowded near tip of 
branchlets; blades somewhat broadly ovate, abruptly 
acuminate with acumen to 15 mm. long, obtuse to 
rounded at base, rarely oblique, serrulate along margins, 
17-28 cm. long, 9-15 cm. wide, chartaceous, in dry state 
dark dirty olive-brown above, grey-green beneath, some- 
what scabrous above but somewhat soft beneath, second- 
ary veins 16-22 pairs, tertiary veins elevated, more 
prominent than lesser venation, strigillose (trichomes 
with barbules) along veins with pustulate epidermis 
above, abundantly and softly stellate (trichomes very 
slender) along and between veins but mixed with un- 
branched multicellular trichomes along major veins with 
somewhat postulate epidermis beneath; petioles 1.5—3 
cm. long, 2-8 mm. in diameter, half-terete, densely pul- 
verulent strigillose to strigose. Inflorescences crowded 
near tip of branchlets, straight, lax and somewhat spread- 
ing, ca. 100-flowered, 18-26 cm. long, 10-17 cm. wide, 
abundantly to densely pulverulent strigose to strigillose 
(trichomes with barbules), primary peduncle 8-11 cm. 
long, bracts triangular to linear, to 6 mm. long. Flow- 
ers 20-25 mm. broad, buds to 8 mm. in diameter, pedi- 
cels 5-12 mm. long, braeteoles minute, subulate, to 2 
mm. long; sepals 5, orbicular, rarely oblong, rounded, 
rarely obtuse, 4-6 mm. long, 4-5 mm. wide, outer 2 
abundantly pulverulent-stellate mixed with unbranched 
multicellular trichomes (with barbules), imbricate one 
abundantly pulverulent-stellate on outer half, densely 
pulverulent-stellate on inner half, inner 2 medially abun- 
dantly pulverulent-stellate, laterally sparingly pulveru- 
lent-stellate, all abundantly to densely appressed-stellate 


[ 269 | 


on upper half, glabrous on lower half within, marginally 
and apically ciliolate to ciliate; petals 5, white, oblong 
to oblong-obovate, 9-11 mm. long, 5-7 mm. wide, sta- 
mens 100-200, filament 3.5-4.5 mm. long, anther 1.5 
mm. long; ovary 5-locular, globose, 5-suleate, glabrous, 
styles 5, 1 mm. long, stigmas simple. Berries not known. 


Distribution: Keuador (Provincia Esmeraldas), river 
bank, alt. 95 m., flowering in December. 


Specimens evamined. Ecuador, Esmeraldas; Parroquia 
de Concepcion, below Playa Rica, Meawia 8488 (F, NY, 
S, U, UC, US-holotype). 


4° Saurauia chaparensis Socjarto sp. nov. 


Species foliis subtus abundanter stellatis, inflorescentiis 
laxis, floribus circiter 15 mm. latis, staminibus plus quam 
100.—Holotypus: Stembach 8920 (GH). 

Shrubs?/: copiously pubescent. Branchlets moderately 
stout, terete, prominently scarred, abundantly strigose 
often with barbulate trichomes to glabrescent. Leaves 
crowded near tip of branchlets ; blades obovate to oblong- 
obovate, blunt to very shortly acuminate at apex, cuneate 
at base, rarely oblique, serrulate with very fine serrula- 
tions along margins, 10-18 em. long, 3-6 cm. wide, 
chartaceous, in dry state sooty above, olive-brown be- 
neath, distinctly scabrous above and beneath, secondary 
veins 18-22 pairs, tertiary veins elevated, more promi- 
nent than lesser venation, sparingly clustered and strigil- 
lose along and between veins with pustulate epidermis 
above, abundantly stellate to rarely tufted along and be- 
tween veins (often mixed with barbulate trichomes along 
major veins) beneath, villous-barbate at axils of second- 
ary veins beneath; petioles 2-8 em. long, 1.5—-2 mm. in 
diameter, terete to half-terete, abundantly tuberculate 
to strigillose with often barbulate trichomes, scabrous. 


[ 270 ] 


Inflorescences crowded near tip of branchlets, straight, 
somewhat loose, 30-80-flowered, 12-20 cm. long, 5-12 
cm. wide, densely pulverulent-tuberculate, primary pe- 
duncle 5-10 cm. long, bracts linear, to 6 mm. long, very 
rarely foliaceous, to 25 mm. long. Flowers ca. 15 mm. 
broad, buds to 6 mm. long, very rarely foliaceous, to 25 
mm. long. Flowers ca. 15 mm. broad, buds to 6 mm. 
in diameter, pedicels 5—-15(—20) mm. long, bracteoles 
triangular, to 2 mm. long; sepals 5, outer 2 oblong- 
obovate, acute, 3.5-4.5 mm. long, 3-3.5 mm. wide, 
densely strigillose (trichomes with barbules), imbricate 
one suborbicular, rounded, 4-5 mm. long, 8.5-4 mm. 
wide, densely strigillose on outer half (trichomes with 
barbules), completely glabrous on inner half, inner 2 
suborbicular to orbicular-oblong, 4-5 mm. long, 8.5—-4.5 
mm. wide, medially densely strigillose (trichomes with 
barbules), laterally completely glabrous, all completely 
glabrous within, marginally subentire, apically irregular- 
ly ciliolate; petals 5, oblong to oblong-obovate, rounded 
to truncate and often incised, 7-9 mm. long, 5-7.5 mm. 
wide; stamens 100-150, filament 2-8 mm. long, anther 
1-2 mm. long; ovary 5-7-locular, globose, 5—7-sulcate, 
glabrous, styles 5-7, obsolete to 4 mm. long, stigmas 
simple to capitate. Berries 5—7-locular, globose, to 5 mm. 
in diameter (immature), 5—7-sulcate. 


Distribution: Bolivia (Departamento Cochabamba), 
forest, alt. 2200-2400 m., flowering in January and 
March. 


Specimens examined. Bolivia, Cochabamba: Chapare, 
Ineachaca, Steinbach 8920 (F, GH-holotype, NY, 5S); 
ibid., Steinbach 9513 (F, G, GH, K, NY, S). 


[ 271 ] 


A7A 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


VoL. 22, No. 8 


CamsBripGk, Massacuuserrs, Drecemper 26, 196 


NATURAL AND ARTIFICIAL HYBRID 
GENERIC NAMES OF ORCHIDS 


SUPPLEMENT IL: 1966-1969 
BY 
LEsLIE A. GARAY AND HERMAN R. SWEET 


Since the publication of the first compendium of ‘‘Nat- 
ural and Artificial Hybrid Generic Names of Orchids, 
1887-1965°’, in the Botanical Museum Leaflets, volume 
21: 141-212, 1966, we have uncovered several previously 
unrecorded names in addition to those which have been 
published for the first time during the intervening period. 
These names, together with corrections and supplemen- 
tary information, are included in this Supplement. Since 
hybrid generic names are not recorded in Index Kewen- 
sis, the first compendium and this supplement provide a 
similar tool for the interested orchidophiles. ‘To increase 
its usefulness, each hybrid generic name is evaluated and 
treated in accordance with the rules set forth in both the 
International Code of Botanical Nomenclature and the 
International Code of Nomenclature for Cultivated 
Plants. Both of these codes derive their authority from 
the International Union of Biological Sciences. 

In this Supplement, as in the first compendium, the 
rule of priority is strictly observed, an action most im- 
portant if stabilization in orchid nomenclature is to be 
achieved. Along this line, we have noted the acceptance 
of some necessary changes by the Orchid Registration 


[ 273 | 


Authority, Royal Horticultural Society, England, such 
as Grammatocymbidium, Phalaerianda, Phragmipa- 
phium, Recchara, Zygorhyncha and Zygostylis, while 
to others, the rule of priority has yet to be applied. For 
example, in 1948, the hybrid generic name Lyonara was 
published with the parentage, 7J'richoglottis * Vanda. 
Since this constituted only a bigeneric hybrid, the use 
of the suffix -ara made the name illegitimate. Subse- 
quently, this was corrected to Trichovanda. In 1959, 
Lyonara was published again with a trigeneric paren- 
tage, Cattleya X Laelia X Schomburghia. This Lyonara 
is also illegitimate, since it represents a later homonym 
as well as a superfluous name. The correct name for this 
trigeneric hybrid 1s Schombolaeliocattleya, originally 
published by The Royal Horticultural Society in its 
Dictionary of Gardening, in 1951, thus antedating the 
later homonym. Nevertheless, we find that Lyonara of 
1959, rather than Schombolaeliocattleya of 1951, is 
maintained in the current listings. Other discrepancies: 
the acceptance of Rhynchovanda (1958) rather than 
Vandachostylis (1985), the acceptance of Brassophron- 
itis (1954) rather than Sophrovola (1895). The former 
hybrid name, involving Rhynchostyls and Vanda, is of 
special concern because Vandachostylis was fully de- 
scribed, illustrated in color, and also supplemented by 
anatomical descriptions and illustrations of the first hy- 
brid, V. Bernardu. At the end of this same treatise, 
another hybrid generic name involving Doritis and 
Phalaenopsis, was mentioned, namely Doritaenopsis, 
but without any description or grex epithet. Notwith- 
standing the lack of description and a legitimate grex 
epithet, Doritaenopsis was and still is accepted by the 
Registration Authority, while Vandachostylis was and 
still is rejected in favor of a later illegitimate name. 

In this Supplement, as in the original compendium, 


[ 274 | 


there has been astrict adherence to the requirements of 
the Codes, especially Articles 40, H. 8 and H. 4 of the 
International Code of Botanical Nomenclature. The in- 
terpretation of these articles was clearly explained in the 
introduction to the first compendium, the correctness of 
which is confirmed by ‘‘An Annotated Glossary of Bo- 
tanical Nomenclature’’, prepared on the authorization 
of the Nomenclature Section of the X. International 
Botanical Congress, in 1964. In this glossary on page 
14, under the term ‘‘Formula’’, we find the following 
statement: ‘‘Condensed formulae formed from parts of 
generic names and applied to intergeneric hybrids are 
treated as ‘generic names’ (Arts. H. 3, H. 4) and are 
subject to the rule of priority and the homonym rule.” 

In a few eases, it was discovered that certain names 
had appeared in publications earlier than those which 
were cited in the original list. These corrections are in- 
cluded in this Supplement in Part I, and, as indicated, 
they replace the earlier entries. ‘The citation of place of 
publication for the following names have been corrected: 
Grammatocymbidium, Gymnaplatanthera, Spatho- 
phaius, Symphodontioda, Symphodontoglossum, 
Symphyglossonia and Zygostylis. 

Hybrids reported for the first time for previously pub- 
lished hybrid generic names include, Grammatocym- 
bidium, Renaglottis and Renancentrum. 

Finally, attention is called to the arrangement of 
parent genera in Part II of this Supplement. The ge- 
neric names involved in the makeup of a given hybrid 
genus appear in alphabetical sequence only once under 
each name. 


m— 
nN 
=I 
a, 
wf 

Lo 


Part I 
List of hybrid generic names 


Aeridocentrum in Orch. Rev. 75: February, 1967 
Aerides ~ Ascocentrum 


Ist hybr.: 4. Luke Nok 


Parentage: dAerides flabellatum 


\ 


< Ascocentrum curvifolium 


Ansidium in Orch. Rev. 74: May, 1966 

Ansellia ~ Cymbidium 

Ist hybr.: 4. Bess Waldon 

Parentage: Ansellia africana *. Cymbidium Dunster Castle 


Arachnostylis in Orch. Rev. 74: April, 1966 

Arachnis * Rhynchostylis 

Ist hybr.: 4. Chorchalood 

Parentage: Arachnis Hookeriana * Rhynchostylis gigantea 


Asconopsis in Orch. Rev. 76: February, 1968 
Ascocentrum “ Phalaenopsis 
Ist hybr.: 4d. Mini-Coral 


Parentage: Phalaenopsis Schilleriana 


* 


< Ascocentrum miniatum 


Ascorachnis in Orch. Rev. 75: August, 1967 
Arachnis * Ascocentrum 

Ist hybr.: 4. George Neo 

Parentage: Arachnis Ishbel  Ascocentrum miniatum 


Ascovandoritis in Orch. Rev. 77: September, 1969 
Ascocentrum ™“ Doritis ~ Vanda 

Ist hybr.: 4. Sonnhild Kitts 

Parentage: Doritis pulcherrima “ Ascocenda Red Gem 


Barlaceras in Riviera Scientif. 11: 62, 1924 

Observation: Add Syn.: Barliaceras in Ciferri and Giacomini, No- 
mencl. Fl. Ital. pt. 1: 167, 1950. 

Barliaceras in Ciferri and Giacomini, Nomencl. FI. Ital. pt. 1: 167 

1950 


Observation: See Barlaceras. 


?. 


Bateostylis in Orch. Rev. 75: November, 1967 
Batemannia  Otostylis 
Ist hybr.: B. Silver Star 


Parentage: Batemannia Colleyi  Otostylis brachystalix 


[ 276 ] 


Bifrinlaria in Lager and Hurrell Seedling Listing no. 866-S: p. 1, 
September, 1966. 

Bifrenaria * Mazillaria 

Ist hybr. : unnamed 

Parentage: Mazillaria Sanderiana * Byfrenaria Harrisoniae 

Bloomara in Orch. Rev. 74: June, 1966 

Broughtonia “~ Laeliopsis << Tetramicra 

Ist hybr.: B. Jim 

Parentage: Jetramicra canaliculata ~ Broughtopsis Kingston (as Lio- 


ponia Kingston) 


Burkillara in Orch. Rev. 75: December, 1967 

Aerides ~ Arachnis ~ Vanda 

Ist hybr.: B. Margaret Ede 

Parentage: Aeridachnis Bogor * /’anda Cooperi (in Orch. Rev. 76: 
June, 1968) 

Observation: For the hybrid Burkillara Henry see Wrefordara. 


Carterara in Orch. Rev. 77: January, 1969 

Aerides * Renanthera  Vandopsis 

Ist hybr.: C. Evening Glow 

Parentage: Renanopsis Lena Rowold  derides Lawrenceae 


Catamodes in Orch. Rev. 75: 205, 1967 

Catasetum “ Mormodes 

Ist hybr.: C. Brown Derby 

Parentage: Catasetum Warscewicsii * Mormodes atropurpurea 


Catanoches in Orch. Rev. 75: 205, 1967 

Catasetum * Cycnoches 

Ist hybr.: C. Green Beret 

Parentage: Catasetum Warscewiczii  Cycnoches ventricosum 


Cattleyopsisgoa in Orch. Rev. 75: November, 1967 
Cattleyopsis ~ Domingoa 

Ist hybr.: C. Little Fellow 

Parentage: Domingoa hymenodes  Cattleyopsis Ortgiesiana 


Cattleyopsistonia in Orch. Rev. 74: July, 1966 
Broughtonia  Cattleyopsis 

lst hybr.: C. Leona 

Parentage: Cattleyopsis Ortgiesiana * Broughtonia sanguinea 
Cephalophrys in Orchid World 2: 114, 1912 

Cephalanthera * Ophrys 


[ 277 ] 


Ist. hybr.: C. integra 
Parentage: Ophrys apifera ~ Cephalanthera rubra 


Christieara in Orch. Rev. 77: November, 1969 
derides * Ascocentrum X Vanda 
Ist hybr.: not yet reported 


Observation: For the hybrid Christieara Mem. Lillian Arnold see 
Reinikkaara. 


Cochlenia in Orch. Rev. 75: November, 1967 

Cochleanthes  Stenia 

Ist hybr.: C. Bryn Mawr 

Parentage: Stenia pallida » Cochleanthes discolor 

Coeloplatanthera in Ciferri and Giacomini, Nomencl. F]. Ital. pt. 1: 
169, 1950 

Coeloglossum »~ Platanthera 

Ist hybr.: C. Brueggeri 


Parentage: Coeloglossum viride  Platanthera chlorantha 


Cymbiphyllum in Orch. Rev. 75: March, 1967 
Observation: See Grammatocymbidium. 


Dactylanthera in Willis, Dict. Fl. Plants and Ferns ed. 7; 327, 1966 
Observation: See Rhizanthera. 


Dactyleucorchis in Ann. Univ. EKétvés, Budapest 8: 319, 1966 
Dactylorhiza ~ Leucorchis 

Ist hybr.: D. Bruniana (as Orchis Bruniana) 

Parentage: Leucorchis albida * Dactylorhiza maculata 


Dactylella in Ann. Univ. Eétvés, Budapest 8: 318, 1966 
Observation: See Dactylitella. 


Dactylitella in Watsonia 6: 132, 1965 

Observation: Add Syn.: Dactylella in Ann. Univ. Eétvés, Budapest 
8: 318, 1966 

Dactyloceras nom. hybr. gen. nov. 

Aceras X Dactylorhiza 

Ist hybr.: D. helvetica comb. nov. (Basionym: Orchiaceras helvetica 
Cif. & Giae., Nomencl. Fl]. Ital, pt. 1: 167, 1950) 

Parentage: Aceras anthropophora * Dactylorhiza latifolia 


Dactylodenia in Bot. Mus. Leafl. Harvard Univ. 21: 157, 1966 

Dactylorhiza * Gymnadenia 

Ist hybr.: D. Heinzeliana comb. nov. (Basionym: Orchis Heinzeliana 
Reich. in Verh. K.K. zool. bot. Ges. 26: 464, 1876) 


[ 278 ] 


Parentage: Dactylorhiza maculata * Gymnadenia conopea 
Syn. : Dactylogymnadenia in Ann. Univ. Eétvés, Budapest 8: 318, 
1966 


Dactylogymnadenia in Ann. Univ. Eétvés, Budapest 8: 318, 1966 
Observation: See Dactylodenia. 


Degarmoara in Orch. Rev. 76: February, 1968 
Brassia X Miltonia X Odontoglossum 

Ist hybr.: D. Agnes 

Parentage: Odontonia Debutante X Brassia antherotes 


DeWolfara nom. hybr. gen. nov. 

Renanthera X Ascoglossum X Ascocentrum * Euanthe X Vanda 

Ist hybr.: D. Cassino (as Shigeuraara Cassino) 

Parentage: Renanthoglossum Red Delight X Schlechterara Meda 
Arnold 


Dialaeliopsis in Orch. Rev. 74: November, 1966 
Diacrium X Laeliopsis 
Ist hybr.: D. Tobago 


Parentage: Diacrium bicornutum  Laeliopsis domingensis 


Dillonara in Orch. Rev. 74: December, 1966 

Epidendrum X Laelia * Schomburgkia 

Ist. hybr.: D. Bronze Kahili 

Parentage: Schombolaelia Maunalani  Hpidendrum diurnum 


Doricentrum in Orch. Rev. 77: October, 1969 
Doritis * Ascocentrum 
Ist. hybr.: D. Merrilee Wallbrunn 


Parentage: Doritis pulcherrima X Ascocentrum curvifolium 


Doriella in Orch. Rev. 74: May, 1966 

Doritis X Kingiella 

Ist hybr.: D. Tiny 

Parentage: Doritis pulcherrima X Kingiella philippinensis 


Doriellaopsis in Orch. Rev. 76: December, 1968 

Doritis X Kingiella X Phalaenopsis 

Ist hybr.: D. Burma 

Parentage: Doritaeniopsis Purple Gem X Kingiella taenialis 


Encyclipedium Dillon in Amer. Orch. Soc. Bull. 38: 676, 1969 
Encyclia * Cypripedium 
Ist hybr.: not yet reported 


Ernestara in Orch. Rev. 76: December, 1968 

Phalaenopsis X Renanthera X Vandopsis 

Ist hybr.: KE. Helga Reuter 

Parentage: Renanopsis Cape Sable X Phalaenopsis Dos Pueblos 


Garayara in Amer. Orch. Soc. Bull. 38: 676, 1969 

Arachnis X Paraphalaenopsis * Vandopsis 

Ist hybr.: G. Lee Kim Hong (as Laycockara Lee Kim Hong) 
Parentage: Pararachnis Lee Siew Chin X Vandopsis lissochiloides 


Gauntlettara in Orch. Rev. 74: April, 1966 

Broughtonia X Cattleyopsis < Laeliopsis 

Ist hybr.: G. Noel 

Parentage: Cattleyopsis Ortgiesiana X Broughtopsis Kingston (as 
Lioponia Kingston) 


Giddingsara in Amer. Orch. Soc. Bull. 38: 676, 1969 

Ascocentrum X Euanthe X Renanthera X Vanda X Vandopsis 

Ist hybr.: G. Sapphire (as Onoara Sapphire) 

Parentage: Schlechterara Meda Arnold X Renanopsis Lena Rowold 


Gilmourara nom. hybr. gen. nov. 

Aerides X Arachnis X Ascocentrum X Euanthe X Vanda 
Ist hybr.: G. Gracia (as Lewisara Gracia) 

Parentage: Aeridachnis Bogor X Schlechterara Ophelia 


Grammatocymbidium in Orch. Rev. 30: 338, 1922 
Cymbidium X Grammatophyllum 
ist hybr.: G. Emil Anderson (as Cymbiphyllum Emil Anderson) 
Parentage: Cymbidium pendulum X Grammatophyllum Measuresianum 
Observation: This replaces the earlier entry. 

Syn.: Cymbiphyllum in Orch. Rev. 75: March, 1967 


Greatwoodara nom. hybr. gen. nov. 

Ascocentrum X Euanthe X Renanthera * Vanda 

Ist hybr.: G. William Doi (as Kagawara William Doi) 
Parentage: Renanthera Kilauea X Schlechterara Meda Arnold 


Gymleucorchis in Lond. Cat. Brit. Pl., ed. 11; 438, 1925 
Gymnadenia * Leucorchis 
Ist hybr.: G. Schweinfurthii (as Gymnadenia Schweinfurthii, 1865) 
Parentage: Gymnadenia conopea X Leucorchis albida 
Ill.: in Verh. Zool.-Bot. Gesellsch. Wien 15: t. 5, f. 15-16, 1865 
Observation: This replaces the earlier entry of Gymnorchis. 

Syn.: Gymnabicchia in Camus, Monogr. Orch. Europ. 315, 1908 


[ 280 | 


Leucadenia in Fedde Rep. 16: 290, 1920, not Leucadenia 
Klotzsch, 1864 

Gymnorchis in Dostal, Fl. Czechosl. (Kvetena CSR) ed. 2, 2101, 
1950 


Gymnabicchia in Camus, Monogr. Orch. Europ. 315, 1908 
Observation: See Gymleucorchis. This replaces the earlier entry. 


Gymnorchis in Dostal, Fl. Czechosl. (Kvetena CSR) ed. 2, 2101, 
1950 
Observation: See Gymleucorchis. This replaces the earlier entry. 


Gymnotraunsteinera in Ciferri and Giacomini, Nomencl. Fl. Ital. 
pt. 1: 171, 1950 

Gymnadenia X Traunsteinera 

Ist hybr.: G. Vallesiaca (as Orchis Vallesiaca) 


Parentage: Gymnadenia conopea X Traunsteinera globosa 


Gymnaplatanthera in Lambert, Notes Orch. Hybr. 9, 1907 
Gymnadenia X Platanthera 
Ist hybr.: G. Chodati (as Gymnadenia Chodati) 
Parentage: Gymnadenia conopea X Platanthera bifolia 
Ill.: in Fedde Rep. Sonderbeih. A. 5: t. 434, f. 1, 1939 
Observation: This replaces the earlier entry of Gymnplatanthera. 
Syn.: Gymnplatanthera in Camus, Monogr. Orch. Europ, 337, 
1908 
Gymplatanthera in Winchester Coll. N.H. Rep. 33, 1911 


Gymnplatanthera in Camus, Monogr. Orch. Europ. 337, 1908 
Observation: See Gymnaplatanthera. This replaces the earlier entry. 
Gymplatanthera in Winchester Coll. N.H. Rep. 33, 1911 
Observation: See Gymnaplatanthera. 

Hawkesara in Orch. Rev. 76: November, 1968 

Cattleya X Cattleyopsis * Epidendrum 

Ist hybr.: H. Alex 

Parentage: Epicattleya Frances Dyer X Cattleyopsis Ori giestana 
Herbertara in Orch. Rev. 76: December, 1968 

Cattleya X Laelia X Schomburgkia X Sophronitis 

ist hybr.: H, Thelma 

Parentage: Schombocattleya Harry Dunn X Sophrolaeliocattleya 


Radians 


Hermibicchia in Camus, Monogr. Orch. Europ. 312, 1908 
Observation: See Herminorchis. This replaces the earlier entry. 


[ 281 ] 


Hermileucorchis in Ciferri and Giacomini, Nomencl. Fl. Ital. pt. 1: 

169, 1950 

Observation: See Herminorchis. 

Herminorchis in Fourn., Quatre Fl. Fr. 201, 1935 

Herminium X Leucorchis 

Ist hybr.: H. Aschersoniana (as Gymnadenia Aschersoniana, 1888) 
Parentage: Leucorchis albida % Herminium monorchis 

Observation: This replaces the earlier entry of Leucerminium and 

Herminiorchis. 

Syn.: Hermibicchia in Camus, Monogr. Orch. Europ. 312, 1908 
Leucerminium in Gartenfl. 85: 258, 19386 
Hermileucorchis in Ciferri and Giacomini, Nomencl. F1]. 

Ital. pt. 1: 169, 1950 
Huntara nom. hybr. gen. nov. 
Arachnis X Euanthe X Renanthera X Vanda * Vandopsis 
Ist hybr.: H. Teoh Cheng Swee (as Teohara Teoh Cheng Swee) 
Parentage: Arandanthe Kian Kee X Renanopsis Lena Rowold 


Huntleanthes in Orch. Rev. 74: January, 1966 
Cochleanthes * Huntleya 
Ist hybr.: H. Narberth 


Parentage: Coch/eanthes discolor X Huntleya Burtii (meleagris) 


Ionettia in Orch. Rev. 76: May, 1968 
Comparettia X lonopsis 


Ist hybr.: /. Rose Petal 


Parentage: Comparettia falcata X lonopsis paniculata 


Ioncidium in Orch. Rev. 76: December, 1968 

Observation: See Ionocidium including the hybrid, Ioncidium Little 
Bit. 

Ionocidium in Orch. Rev. 76: May, 1968 

fonopsis X Oncidium 

Ist hybr.: /. Ressie Toy 

Parentage: /onopsis paniculata X Oncidium pulchellum 
Syn.: Ioncidium in Orch. Rev. 76: December, 1968 


Jacquinparis Dillon in Amer. Orch. Soc. Bull. 38: 676, 1969 
Jacquiniella X Liparis 
Ist hybr.: not yet reported 


Kagawara in Orch. Rev. 76: June, 1968 
Ascocentrum X Renanthera X Vanda 
Ist hybr.: A. Firebird (Orch. Rev. 77: March, 1969) 


{ 282 ] 


Parentage: Renanthera Storiei X Ascocenda Red Gem 

Observation: for the hybrid Kagawara William Doi see Greatwood- 
ara. 

Kamemotoara nom. hybr. gen. nov. 

Aerides X Euanthe X Rhynchostylis * Vanda 

Ist hybr.: A. Porchina Blue (as Perreiraara Porchina Blue) 

Parentage: Rhynchovandanthe Blue Angel X derides mitrata 


Laeliocatonia in Orch. Rev. 75: March, 1967 

Broughtonia X Cattleya X Laelia 

Ist hybr.: L. Betty Holley 

Parentage: Laeliocattleya Bright Night Xx Broughtonia sanguinea 


Laeliopleya in Orch. Rev. 74: March, 1966 

Cattleya * Laeliopsis 

Ist hybr.: L. Orange Glow 

Parentage: Cattleya aurantiaca X Laeliopsis domingensis 


Laycockara in Orch. Rev. 74: August, 1966 

Arachnis X Phalaenopsis X Vandopsis 

Ist hybr. : not yet reported 

Observation: For the hybrids Laycockara Lee Kim Hong and L. 


lan Trevor, see Garayara. 


Leucadenia in Fedde Rep. 16: 290, 1920 
Observation: Not Leucadenia Klotzsch, 1864. See Gymleucorchis. 
This replaces the earlier entry. 


Leucerminium in Gartenfl. 85: 253, 1936 
Observation: See Herminorchis. This replaces the earlier entry. 


Leucororchis in Ciferri and Giacomini, Nomencl. Fl]. Ital. pt. 1: 
170, 1950 

Leucorchis X Orchis 

Ist hybr.: not yet reported 

Observation: For the hybrids Leucorochis Bruniana and L. albucina 
see Dactyleucorchis. 


Lewisara in Orch. Rev. 76: February, 1968 

Aerides X Arachnis X Ascocentrum X Vanda 

Ist hybr.: L. Max 

Parentage: Aeridachnis Bogor X Ascocenda Charm 
Observation: For the hybrid Lewisara Gracia see Gilmourara. 


Loroglorchis in Journ. Bot. Fr. 6: 110, 1892 


Observation: This replaces the earlier entry. See Orchimantoglos- 
sum. 


[283 ] 


37 


Lymanara in Orch. Rev. 75: August, 196 
Aerides X Arachnis * Renanthera 

Ist hybr.: L. Mary Ann 

Parentage: Aeridachnis Bogor X Renanthera Storiei 


Macradesa in Orch. Rev. 76: December, 1968 
Gomesa * Macradenia 
Ist hybr.: M. Brown Baby 


Parentage: Gomesa recurva X Macradenia brassavolae 


Maxillacaste in La Sem. Hortic. I: 350, 1897 
Mawillaria X Lycaste 
Ist hybr.: not yet reported 


Mizutara in Orch. Rev. 74: August, 1966 

Cattleya X Diacrium X Schomburgkia 

Ist hybr.: M. Pink Kahili 

Parentage: Schombocattleya Diamond Head & Diacrium bicornutum 


Mokara in Orch. Rev. 77: December, 1969 
Arachnis * Ascocentrum X Vanda 

Ist hybr.: M. Wai Liang 

Parentage; Arachnis Ishbel X Ascocenda Red Gem 


Moscosoara in Orch. Rev. 77: September, 1969 
Broughtonia X Epidendrum X Laeliopsis 

Ist hybr.: M. Santo Domingo 

Parentage: Broughtopsis Kingston * Epidendrum olivaceum 


Myrmecocattleya in Orch. Rev. 28: 50, 1920 
Observation: See Schombocattleya. 


Myrmecolaelia in Orch. Rev. 28: 50, 1920 

Observation: See Schombolaelia. 

Nobleara in Orch. Rev. 77: September, 1969 

Aerides X Renanthera X Vanda 

Ist hybr.: N. Royal Monarch 

Parentage: Renantanda Jukichi Murata X Aerides odorata 


Oncidenia in Orch. Rev. 74: February, 1966 

Macradenia X Oncidium 

Ist hybr.: O. Helen 

Parentage: Oncidium Helen Brown X Macradenia brassavolae 


Oncidpilia in Orch. Rev. 74: February, 1966 
Oncidium X Trichopilia 
Ist hybr.: O. Don Carlos 

[ 284 ] 


Parentage: Oncidium Papilio * Trichopilia coccinea 


Onoara in Orch. Rev. 75: August, 1967 

Ascocentrum X Renanthera X Vanda X Vandopsis 

ist hybr.: not yet reported 

Observation: For the hybrid Onoara Sapphire see Giddingsara. 


Orchidatyla in Watsonia 6: 133, 1965 
Observation: Syn.: Orchidactylorhiza in Ann. Univ. Eétvés, Buda- 
pest 8: 315, 1966 


Orchidactylorhiza in Ann. Univ. Eétvés, Budapest 8: 315, 1966 
Observation: See Orchidactyla. 


Orchimantoglossum in Aschers. & Graebn., Syn. 3: 799, 1907 
Orchis X Himantoglossum 
Ist hybr.: O. Lacasei 
Parentage: Orchis simia X Himantoglossum hircinium 
Observation: Because Himantoglossum is conserved over Loroglossum 
this replaces the earlier entry. 
Syn.: Loroglorchis in Journ. Bot. Fr. 6: 110, 1893 


Osmentara in Orch. Rev. 74: June, 1966 

Broughtonia X Cattleya X Laeliopsis 

Ist hybr.: O. Bill 

Parentage: Cattleya R. Prowe X Broughtopsis Kingston (as Lioponia 
Kingston) 

Otonisia in Orch. Rev. 77: January, 1969 

Aganisia X Otostylis 

Ist hybr.: O. Broadway 


Parentage: Otostylis brachystalix X Aganisia pulchella 


Pattoniheadia Dillon in Amer. Orch. Soc. Bull. 38: 676, 1969 
Pattonia X Bromheadia 
Ist hybr. : not yet reported 


~ 


Perreiraara in Orch. Rev. 77: September, 1969 

Aerides X Rhynchostylis * Vanda 

Ist hybr.: not yet reported 

Observation: For the hybrid Perreiraara Porchina Blue see Kame- 
motoara. 

Pterocottia Dillon in Amer. Orch. Soc. Bull. 38: 676, 1969 

Plerostylis X Prescottia 

Ist hybr.: not yet reported 


Reinikkaara nom. hybr. gen. nov. 


[ 285 | 


Aerides X Ascocentrum X Euanthe X Vanda 

Ist hybr.: R. Mem. Lillian Arnold (as Christieara Mem. Lillian 
Arnold) 

Parentage: Aerides Lawrenceae X Schlechterara Meda Arnold 


Renaglottis in Bull. Pac. Orch. Soc. Hawaii 14: 85, 1957 
Renanthera X Trichoglottis 

Ist hybr.: R. Lone Warrior (in Orch. Rev. 74: November, 1966) 
Parentage: Renanthera Storiei * Trichoglottis fasciata 


Renancentrum in Orch. Rev. 70: September, 1962 

Ascocentrum X Renanthera 

Ist hybr.: R. Yap Sin Yee (in Orch. Rev. 73: May, 1965) 
Parentage: Ascocentrum curvifolium X Renanthera Brookie Chandler 
Observation: For the hybrid Renancentrum Curvionica see Ascorella. 


Renanthoceras nom. hybr. gen. nov. 
Pteroceras X Renanthera 
Ist hybr.: FR. Kona (as Sarcothera Kona) 


Parentage: Pteroceras pallidus X Renanthera monachica 


Renanthopsis in Bull. Soc. Hort. Fr. ser. 5, 4: 342, 1931 
Observation: This replaces the earlier entry. 


Restesia Dillon in Amer. Orch. Soc. Bull. 38: 676, 1969 
Restrepia X Orleanesia 
Ist hybr.: not yet reported 


Rhizanthera in Watsonia 6: 138, 1965 

Observation: Add Syn.: Dactylanthera in Willis, Dict. Fl. Plants 
and Ferns, ed. 7: 327, 1966 

Rhyndoropsis in Orch. Rev. 74: October, 1966 

Doritis X Phalaenopsis * Rhynchostylis 

Ist hybr.: FR. Florida 

Parentage: Doritaenopsis Dorette < Rhynchostylis retusa 


Rodrenia in Amer. Orch. Soc. Bull. 31: 357, 1962 
Observation: See Rodridenia. 


Rodridenia in Orch. Rev. 70: April, 1962 

Observation: Add Syn.: Rodrenia in Amer. Orch. Soc. Bull. 31: 
357, 1962 

Rodriopsis in Orch. Rev. 77: September, 1969 

Rodriguesia X lonopsis 

Ist hybr.: FR. Edwardine Klemm 

Parentage: Rodriguesia secunda  Ionopsis paniculata 


[ 286 ] 


Rumrillara in Orch. Rev. 77: September, 1969 

Ascocentrum X Neofinetia X Rhynchostylis 

Ist hybr.: R. Rosyleen 

Parentage: Rhynchocentrum Lilac Blossom X Ascofinetia Peaches 


Sarcorhiza in Orch. Rev. 74: January, 1966 

thinerrhiza X Sarcochilus 

Ist hybr.: S. Doroty 

Parentage: Sarcochilus Hartmanit * Rhinerrhiza divitiflora 


Sarcothera in Orch. Rev. 62: 92, 1954 

Renanthera X Sarcochilus 

Ist hybr.: not yet reported 

Observation: For the hybrid Sarcothera Kona see Renanthoceras. 


This replaces the earlier entry. 


Schombocattleya in Orch. Rev. 13: 245, 1905 
Observation: Add Syn. : Schomocattleya in Orch. Rev. 28: 50, 1920 
Myrmecocattleya in Orch. Rev.28 : 50, 1920 


Schombolaelia in Orch. Rev. 21: 254, 1913 
Observation: Add Syn.: Myrmecolaelia in Orch. Rev. 28: 50, 1920 


Schombonitis in Orch. Rev. 28: 50, 1920 

Schomburgkia X Sophronitis 

Ist hybr.: S. Stella 

Parentage: Schomburgkia superbiens * Sophronitis grandiflora 


Schomocattleya in Orch. Rev. 28: 50, 1920 
Observation: See Schombocattleya. 


Selenipanthes Dillon in Amer. Orch. Soc. Bull. 58: 676, 1969 
Selenipedium X Lepanthes 
Ist hybr. : not yet reported 


Serapirhiza in Jahresb. Naturwiss. Ver. Wuppertal, Heft 21/22: 
103, 1968 
Serapias * Dactylorhiza 
Ist hybr. : S. Sambucino-lingua (Barla) comb. nov. 
(Basionym: Orchis sambucina var. sambucino-lingua Barla, Icon. 
Orch. 60, 1868) 
Parentage: Serapias Lingua X Dactylorhiza sambucina 


Shigeuraara in Orch. Rev. 77: November, 1969 

Renanthera X Ascoglossum X Ascocentrum * Vanda 

Ist hybr.: not yet reported 

Observation: For the bybrid Shigeuraara Cassino see DeWolfara. 


[ 287 ] 


Spathophaius in Orchid Weekly 1: 268, 1959 
Observation: This replaces the earlier entry. 


Stanfieldara in Orch. Rev. 77: November, 1969 
Sophronitis < Laelia * Epidendrum 

Ist hybr.: S. Will Bates 

Parentage: Sophrolaelia Psyche X Epidendrum vitellinum 


Sweetara in Amer. Orch. Soc. Bull. 38: 676, 1969 

Paraphalaenopsis * Rhynchostylis X Vanda 

ist hybr.: S. Oi Yee (as Yapara Oi Yee) 

Parentage: Paravanda Suavei * Vandachostylis Tan Geat Leng (as 
Rhynchovanda ‘T'an Geat Leng) 


Symphodontioda in Orchid Weekly 4: 121, 1963 
Observation: This replaces the earlier entry. 


Symphodontoglossum in Orchid Weekly 4: 121, 1963 
Observation: This replaces the earlier entry. 


Symphyglossonia in Orchid Weekly 4: 121, 19638 
Observation: This replaces the earlier entry. 


Teohara in Orch. Rev. 76: February, 1968 

Arachnis X Renanthera X Vanda X Vandopsis 

Ist hybr.: not yet reported 

Observation: For the hybrid Teohara Teo Cheng Swee see Huntara 


Vanglossum in Orch. Rev. 77: March, 1969 
Ascoglossum X Vanda 

Ist hybr.: V’, Oriental Jewel 

Parentage: Vanda Merrillii X Ascoglossum caloplerum 


Withnerara in Orch. Rev. 74: November, 1966 
Aspasia X Miltonia X Odontoglossum X Oncidium 

Ist hybr.: JV. Moon Glow 

Parentage: Oncidasia Starlight X Odontonia Wonder 


Wrefordara in Amer. Orch. Soc. Bull. 38: 676, 1969 
Aerides X Arachnis X Euanthe X Vanda 

Ist hybr.: W. Henry (as Burkillara Henry ) 

Parentage: Aeridachnis Bogor X Vandanthe Ellen Noa 


Yapara in Orch. Rev. 74: November, 1966 

Phalaenopsis X Rhynchostylis X Vanda 

Ist hybr.: not yet reported 

Observation: For the hybrid Yapara Oi Yee see Sweetara. 


[ 288 ] 


Zygomena in Die Natuerl. Pflanzenfam. Erg.-heft Il, 92, 1908 
Zygopetalum X Zygosepalum (Menadenium) 

Ist hybr.: Z. Roeblingiana (as Zygopetalum Roeblingianum) 
Parentage: Zygopetalum maxillare X Zygosepalum labiosum 

Ill.: in Gard. Chron. ser. 3, 34: 227, f. 98, 1903 
Observation: This replaces the former entry. 


Zygostylis in Handbook Orch. Nomencl. and Reg., p. 47, 1965 
Observation: See Orch. Rev. 74: April, 1966. This replaces the 
earlier entry. 


Parr II 
Parentage index to hybrid generic names 


Parent genera Hybrid genus 


Aceras X Dactylorhiza = Dactyloceras 


Aerides X Arachnis X Ascocentrum X Euanthe 


Xx Vanda = Gilmourara 
Aerides X Arachnis X Ascocentrum X Vanda = Lewisara 
Aerides X Arachnis X Euanthes X Vanda = Wrefordara 
Aerides X Arachnis X Renanthera = Lymanara 
Aerides X Arachnis X Vanda = Burkillara 
Aerides X Ascocentrum = Aeridocentrum 
Aerides X Ascocentrum X Euanthe X Vanda = Reinikkaara 
Aerides X Ascocentrum X Vanda = Christieara 
Aerides X Euanthe X Rhynchostylis X Vanda = Kamemotoara 
Aerides X Renanthera X Vanda = Nobleara 
Aerides X Renanthera X Vandopsis = Carterara 
Aerides X Rhynchostylis X Vanda = Perreiraara 
Aganisia X Otostylis = Otonisia 
Ansellia X Cymbidium = Ansidium 
Arachnis X Aerides X Ascocentrum X Euanthe 

X Vanda = Gilmourara 
Arachnis X Aerides X Ascocentrum X Vanda = Lewisara 
Arachnis X Aerides X Euanthe X Vanda = Wrefordara 
Arachnis X Aerides X Renanthera = Lymanara 
Arachnis X Aerides X Vanda = Burkillara 
Arachnis X Ascocentrum = Ascorachnis 
Arachnis X Ascocentrum X Vanda = Mokara 


[ 289 | 


Arachnis X Kuanthe X Renanthera X Vanda 


X Vandopsis = Huntara 
Arachnis X Paraphalaenopsis X Vandopsis = Garayara 
Arachnis < Phalaenopsis X Vandopsis = Laycockara 
Arachnis X Renanthera X Vanda X Vandopsis = Teohara 
Arachnis X Rhynchostylis = Arachnostylis 
Ascocentrum X Aerides = Aeridocentrum 
Ascocentrum X Aerides X Arachnis X Euanthe 

Vanda = Gilmourara 
Ascocentrum X Aerides X Arachnis X Vanda = Lewisara 
Ascocentrum X Aerides X EKuanthe X Vanda = Reinikkaara 
Ascocentrum X Aerides X Vanda = Christieara 
Ascocentrum X Arachnis = Ascorachnis 
Ascocentrum X Arachnis X Vanda = Mokara 


Ascocentrum X Ascoglossum X Euanthe X Renan- 
thera X Vanda = DeWolfara 


Ascocentrum X Ascoglossum X Renanthera X Vanda = Shigeuraara 


Ascocentrum X Doritis = Doricentrum 
Ascocentrum X Doritis X Vanda = Ascovandoritis 
Ascocentrum X Euanthes X Renanthera X Vanda = Greatwoodara 
Ascocentrum X Euanthe X Renanthera X Vanda 

*Vandopsis = Giddingsara 
Ascocentrum X Neofinetia X Rhynchosty lis = Rumrillara 
Ascocentrum X Phalaenopsis = Asconopsis 
Ascocentrum X Renanthera = Renancentrum 
Ascocentrum X Renanthera X Vanda = Kagawara 
Ascocentrum X Renanthera X Vanda X Vandopsis == Onoara 


Ascoglossum X Ascocentrum X Renanthera X Vanda = DeWolfara 
Ascoglossum X Vanda = Vanglossum 


Ascoglossum X Ascocentrum X Renanthera X Vanda = Shigeuraara 


Aspasia X Miltonia X Odontoglossum X Oncidium = Withnerara 
Batemannia X Otosty lis = Bateostylis 
Bifrenaria X Maxillaria = Bifrinlaria 
Brassia X Miltonia X Odontoglossum = Degarmoara 
Bromheadia X Pattonia = Pattoniheadia 
Broughtonia X Cattleya X Laelia = Laeliocatonia 
Broughtonia X Cattleya X Laeliopsis = Osmentara 


[ 290 ] 


Broughtonia X Cattleyopsis 
Broughtonia X Cattleyopsis X Laeliopsis 
Broughtonia X Epidendrum X Laeliopsis 
Broughtonia X Laeliopsis X Tetramicra 


Catasetum X Cycnoches 
Catasetum X Mormodes 


Cattleya X Broughtonia X Laelia 
Cattleya X Broughtonia X Laeliopsis 
Cattleya X Cattleyopsis X Epidendrum 
Cattleya X Diacrium X Schomburgkia 


Cattleya X Laelia X Schomburgkia X Sophronitis 


Cattleya X Laeliopsis 


Cattleyopsis X Broughtonia 
Cattleyopsis X Broughtonia X Laeliopsis 
Cattleyopsis X Cattleya X Epidendrum 
Cattleyopsis X Domingoa 


Cephalanthera X Ophrys 


Cochleanthes X Huntleya 
Cochleanthes XStenia 


Coeloglossum X Platanthera 
Comparettia X lonopsis 
Cyenoches X Catasetum 
Cymbidium X Ansellia 
Cypripedium X Encyclia 


Dactylorhiza X Aceras 
Dactylorhiza X Leucorchis 
Dactylorhiza X Serapias 


Diacrium X Cattleya X Schomburgkia 
Diacrium X Laeliopsis 


Domingoa X Cattleyopsis 


Doritis X Ascocentrum 

Doritis X Ascocentrum X Vanda 

Doritis X Kingiella 

Doritis X Kingiella X Phalaenopsis 
Doritis X Phalaenopsis X Rhynchostylis 


[ 291 ] 


= Cattleyopsistonia 
= Gauntlettara 
= Moscosoara 


= Bloomara 


= Catanoches 


= Catamodes 


= Laeliocatonia 
= Osmentara 

= Hawkesara 

= Mizutara 

= Herbertara 
= Laeliopleya 


= Cattleyopsistonia 
= Gauntlettara 
= Hawkesara 


= Cattleyopsisgoa 
=Cephalophrys 


= Huntleanthes 
=Cochlenia 


= Coeloplatanthera 
= lonettia 

= Catanoches 

= Ansidium 

= Encyclipedium 


= Dactyloceras 
= Dactyleucorchis 
= Serapirhiza 


= Mizutara 


= Dialaeliopsis 
= Cattleyopsisgoa 


= Doricentrum 
= Ascovandoritis 
= Doriella 

= Doriellaopsis 
= Rhyndoropsis 


Encyclia X Cy pripedium = Encyclipedium 


Epidendrum X Broughtonia X Laeliopsis = Moscosoara 
Epidendrum X Cattleya X Cattleyopsis = Hawkesara 
Epidendrum X Laelia X Schomburgkia = Dillonara 
Epidendrum X Laelia X Sophronitis = Stanfieldara 
Kuanthe X Aerides X Arachnis * Ascocentrum 

* Vanda = Gilmourara 
Euanthe X Aerides X Arachnis X Vanda = Wrefordara 
Euanthe X Aerides X Ascocentrum X Vanda = Reinikkaara 
Euanthe X Aerides X Rhynchostylis X Vanda = Kamemotoara 
Euanthe X Arachnis X Renanthera X Vanda 

X Vandopsis = Huntara 


Kuanthe X Ascocentrum X Ascoglossum 
< Renanthera X Vanda = DeWolfara 


Euanthe X Ascocentrum X Renanthera X Vanda = Great woodara 
Euanthe X Ascocentrum X Renanthera 


X Vanda X Vandopsis = Giddingsara 
Gomesa X Macradenia = Macradesa 
Gy mnadenia X Leucorchis = Gymleucorchis 
Gymnadenia X Platanthera = Gymnaplatanthera 
Gymnadenia X Traunsteinera = Gymnotraunsteinera 
Herminium X Leucorchis = Herminorchis 
Himantoglossum X Orchis = Orchimantoglossum 
Huntleya X Cochleanthes = Huntleanthes 
lonopsis X Comparettia = lonettia 
lonopsis X Oncidium = lonocidium 
lonopsis X Rodriguezia = Rodriopsis 
Jacquiniella X Liparis = Jacquinparis 
Kingiella X Doritis = Doriella 
Kingiella X Doritis X Phalaenopsis = Doriellaopsis 
Laelia X Broughtonia X Cattleya = Laeliocatonia 
Laelia X Cattleya X Schomburgkia 

X Sophronitis = Herbertara 
Laelia X Epidendrum X Schomburgkia = Dillonara 
Laelia X Epidendrum X Sophronitis = Stanfieldara 
Laeliopsis X Broughtonia X Cattleya = Osmentara 
Laeliopsis < Broughtonia X Cattleyopsis = Gauntlettara 


[ 292 } 


Laeliopsis X Broughtonia X Epidendrum 
Laeliopsis X Broughtonia X Tetramicra 
Laeliopsis X Cattleya 

Laeliopsis * Diacrium 


Lepanthes X Selenipedium 


Leucorchis X Dactylorhiza 
Leucorchis X Gymnadenia 
Leucorchis X Herminium 
Leucorchis  Orchis 


Liparis X Jacquiniella 
Lycaste X Maxillaria 


Macradenia X Gomesa 
Macradenia X Oncidium 


Maxillaria X Bifrenaria 
Maxillaria < Lycaste 


Miltonia < Aspasia X Odontoglossum 
X Oncidium 
Miltonia X Brassia X Odontoglossum 


Mormodes X Catasetum 
Neofinetia X Ascocentrum X Rhynchostylis 


Odontoglossum X Aspasia X Miltonia 
* Oncidium 
Odontoglossum X Brassia X Miltonia 


Oncidium X Aspasia X Miltonia 
X Odontoglossum 

Oncidium X Ionopsis 

Oncidium X Macradenia 

Oncidium X Trichopilia 

Ophrys X Cephalanthera 


Orchis X Himantoglossum 
Orchis X Leucorchis 


Orleanesia X Restrepia 


Otostylis X Batemannia 
Otostylis X Aganisia 


[293 | 


= Moscosoara 
= Bloomara 

= Laeliopleya 
= Dialaeliopsis 


= Selenipanthes 


= Dactyleucorchis 
= Gymleucorchis 
= Herminorchis 

= Leucororchis 


= Jacquinparis 
= Maxillacaste 


= Macradesa 
= Oncidenia 
= Bifrinlaria 


= Maxillacaste 


= Withnerara 


= Degarmoara 
= Catamodes 


= Rumrillara 


= Withnerara 
= Degarmoara 


= Withnerara 
= Jonocidium 
= Oncidenia 
= Oncidpilia 


=Cephalophrys 


= Orchimantoglossum 
= Leucororchis 
= Restesia 


= Bateostylis 
= Otonisia 


Paraphalaenopsis X Arachnis X Vandopsis = Garayara 


Paraphalaenopsis X Rhynchostylis X Vanda = Sweetara 
Pattonia X Bromheadia = Pattoniheadia 
Phalaenopsis X Arachnis X Vandopsis = Laycockara 
Phalaenopsis X Ascocentrum = Asconopsis 
Phalaenopsis X Doritis X Kingiella = Doriellaopsis 
Phalaenopsis X Doritis X Rhynchostylis = Rhyndoropsis 
Phalaenopsis X Renanthera X Vandopsis = Ernestara 
Phalaenopsis X Rhynchostylis X Vanda = Yapara 
Platanthera X Coeloglossum = Coeloplatanthera 
Platanthera X Gymnadenia = Gymnaplatanthera 
Pteroceras X Renanthera = Renanthoceras 
Renanthera X Aerides X Arachnis = Lymanara 
Renanthera X Aerides X Vanda = Nobleara 
Renanthera X Aerides X Vandopsis =Carterara 
Renanthera X Arachnis X Euanthe X Vanda 

X Vandopsis = Huntara 
Renanthera X Arachnis X Vanda X Vandopsis =Teohara 
Renanthera X Ascocentrum = Renancentrum 
Renanthera X Ascocentrum X Ascoglossum 

X Euanthe X Vanda = DeWolfara 
Renanthera X Ascocentrum X Ascoglossum 

Vanda = Shigeuraara 
Renanthera X Ascocentrum X Euanthe X Vanda =Greatwoodara 
Renanthera X Ascocentrum X Vanda = Kagawara 
Renanthera X Ascocentrum X Vanda 

X Vandopsis = QOnoara 
Renanthera X Phalaenopsis X Vandopsis = Ernestara 
Renanthera X Pteroceras = Renanthoceras 
Renanthera X Sarcochilus = Sarcothera 
Renanthera X Trichoglottis = Renaglottis 
Restrepia X Orleanesia = Restesia 
Rhinerrhiza X Sarcochilus = Sarcorhiza 
Rhynchostylis X Aerides X Kuanthe X Vanda = Kamemotoara 
Rhynchostylis X Aerides X Vanda = Perreiraara 
Rhynchostylis X Arachnis = Arachnostylis 
Rhynchostylis X Ascocentrum X Neofinetia = Rumrillara 


[ 294 ] 


Rhynchostylis X Doritis X Phalaenopsis 
Rhynchostylis X Paraphalaenopsis X Vanda 
Rhynchostylis X Phalaenopsis X Vanda 


Rodriguezia X Ionopsis 


Sarcochilus X Renanthera 
Sarcochilus X Rhinerrhiza 


Schomburgkia X Cattleya X Diacrium 
Schomburgkia X Cattleya X Laelia 

X Sophronitis 
Schomburgkia X Epidendrum X Laelia 
Schomburgkia X Sophronitis 


Selenipedium X Lepanthes 
Serapias X Dactylorhiza 


Sophronitis X Cattleya X Laelia 

X Schomburgkia 
Sophronitis X Epidendrum X Laelia 
Sophronitis X Schomburgkia 


Stenia X Cochleanthes 

Tetramicra X Broughtonia X Laeliopsis 
Traunsteinera X Gymnadenia 
Trichoglottis X Renanthera 
Trichopilia X Oncidium 

Vanda X Aerides X Arachnis 


Vanda X Aerides X Arachnis X Ascocentrum 


Vanda X Aerides X Arachnis X Ascocentrum 
X Euanthe 


Vanda X Aerides X Arachnis X Euanthe 

Vanda X Aerides X Ascocentrum 

Vanda X Aerides X Ascocentrum X Euanthe 

Vanda X Aerides X Euanthe X Rhynchostylis 

Vanda X Aerides X Renanthera 

Vanda X Aerides X Rhynchostylis 

Vanda X Arachnis X Ascocentrum 

Vanda X Arachnis X Euanthe X Renanthera 
X Vandopsis 


r 


[ 295 | 


= Rhyndoropsis 
=Sweetara 
= Yapara 


= Rodriopsis 


= Sarcothera 


= Sarcorhiza 


= Mizutara 


= Herbertara 
= Dillonara 
=Schombonitis 


= Selenipanthes 


= Serapirhiza 


= Herbertara 
= Stanfieldara 
= Schombonitis 


=Cochlenia 

= Bloomara 

= Gymnotraunsteinera 
= Renaglottis 

= Oncidpilia 

= Burkillara 


= Lewisara 


= Gilmourara 

= Wrefordara 

= Christieara 

= Reinikkaara 
= Kamemotoara 
= Nobleara 

= Perreiraara 


= Mokara 


= Huntara 


Vanda X Arachnis X Renanthera X Vandopsis = Teohara 
Vanda X Ascocentrum X Ascoglossum X Euanthe 


X Renanthera = DeWolfara 
Vanda X Ascocentrum X Ascoglossum X Renanthera =Shigeuraara 
Vanda X Ascocentrum X Doritis = Ascovandoritis 
Vanda X Ascocentrum X Euanthe X Renanthera = Greatwoodara 
Vanda X Ascocentrum X Renanthera = Kagawara 
Vanda X Ascocentrum X Renanthera X Vandopsis = Onoara 
Vanda X Ascoglossum = Vanglossum 
Vanda X Paraphalaenopsis X Rhynchostylis = Sweetara 
Vanda X Phalaenopsis X Rhynchosty lis = Yapara 
Vandopsis X Aerides X Renanthera = Carterara 
Vandopsis X Arachnis X Euanthe X Renanthera 

X Vanda = Huntara 
Vandopsis X Arachnis X Paraphalaenopsis = Garayara 
Vandopsis X Arachnis X Phalaenopsis = Laycockara 
Vandopsis X Arachnis X Renanthera X Vanda = Teobara 
Vandopsis X Ascocentrum X Euanthe X Renanthera 

X Vanda = Giddingsara 
Vandopsis X Ascocentrum X Renanthera X Vanda = Onoara 
Vandopsis X Phalaenopsis X Renanthera = Ernestara 
Zy gopetalum X Zy gosepalum = Zygomena 
Zy gosepalum X Zy gopetalum = Zygomena 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


Campripnce, Massacuusetrs, Aprit 30, 1970 VoL. 22, No. 9 


TEOSINTE INTROGRESSION IN THE 
MAIZE OF THE NOBOGAME VALLEY 
BY 
H. Garrison WILKES 


Around Nobogame grows a plant called maizillo, or maizmillo. 
It is more slender than the ordinary corn-plant and the ears are 
very small. It growsamong the corn and has to be weeded out, 
as itinjures the good plants. However, several Mexicans assured 
me that, when cultivated, the ears develop. After three years 
they grow considerably larger and may be used as food. A man 
in Cerro Prieto raises this kind only: others mix it with the 
ordinary corn. I was told that people from the Hot Country 
came to gather it, each taking away about one almud to mix 
with their seed corn. The combination is said to give splendid 
results in fertile soil. 

Lumunottz, Unknown Mexico 
1902 

The Nobogame Valley is an isolated valley in the 
Sierra Madre Occidental of Mexico, and the description 
by Lumholtz written over fifty years ago is an accurate 
account of present day hybridization of teosinte with 
maize. 

This valley, inhabited by Tarahumare Indians and 
Mexicans, is approximately eight miles north of the old 
mining town of Guadalupe y Calvo in the southwestern 
corner of the state of Chihuahua. Most of the valley 
floor is given over to the cultivation of maize, beans and 
squash or grazed, but teosinte does occur and is often 
abundant along the margins of maize fields or in the 
willow thickets bordering the streams. The distribution 


[ 297 ] 


of teosinte is limited to probably not more than 25 square 
miles of the valley between the elevations of 1720 and 
1920 meters. 

The local inhabitants recognize teosinte as distinct 
from maize and call it mazcillo or maiz silvestre. They 
are also familiar with the maize Xteosinte hybrids and 
universally will claim that if the hybrid is cultivated for 
three years it produces maize. Some feel that hybridi- 
zation even improves the maize. 


Factors limiting gene exchange 

Both teosinte and maize are wind-borne, cross-polli- 
nated plants, and the prevention of hybridization with 
subsequent genetic exchange between the two species is 
dependent on (1) the spatial isolation of the two species, 
(2) the seasonal isolation of the two species, (3) the lack 
of fitness of the hybrid, and (4) the types of selection 
operating on the two parental populations. 

The most effective isolating mechanism in most areas 
where teosinte and maize are sympatric has been the 
mean mid-flowering date, with maize flowering 2-5 
weeks ahead of teosinte. Nobogame teosinte is unique 
in that itis the only teosinte population found in a habi- 
tat other than a cultivated field which is not seasonally 
isolated from maize. Maize of the Nobogame Valley is 
a five month type belonging to the race Cristalino de 
Chihuahua which is planted in May and harvested in 
September before the early killing frost. Both teosinte 
and maize reach mid-flowering in August. 


Hybridization at Nobogame 

In all the areas of both Guatemala and Mexico where 
teosinte and maize occur together there is some evidence 
(only a single F; hybrid in some cases) of hybridization, 
but never has the author found such a large number of 
hybrids and clear and unmistakable effect of teosinte in- 


[ 298 | 


Triste Lik 


Maize of the Nobogame Valley. Cristalino de Chihuahua is a poorly 
defined race in Mexico which is typically a dent corn. There is con- 
siderable variation from field to field and very bard flinty kernels are 
frequent in the region where hybridization with teosinte occurs. 2/5 


actual size. 


trogression in maize cobs as exists in Nobogame. The 
abundance of Fy, hybrids is comparable to Chalco, where 
several researchers have studied maize X teosinte hybrids 
(Lopez y Para (1908), Collins (1921), Bukasov (1926, 
1930), Mangelsdorf (1952), and Wilkes (1967)). Unlike 
the region around Chalco, however, teosinte is not lim- 
ited to the cultivated fields. ‘Teosinte also occurs in dense 
stands along the streams and in areas protected from 
grazing on the surrounding hills. 

Although all the fields had been harvested at the time 
of the field work (November), careful investigation indi- 
‘rates that maize Xteosinte hybrids are present in the 
fields proper, but not as abundant as they are on the 
margin of the maize fields or in the willow thickets along 
the stream. When maize X teosinte hybrids are encoun- 
tered in the maize fields of Mexico, it is usually pre- 
sumed that they are from teosinte fruit-cases containing 
hybrid seed which were naturally disseminated in the 
tield. In Nobogame, the female parent of the teosinte x 
maize hybrids is sometimes maize and thus the hybrids 
are often planted in the field. 


Teosinte introgression 


The evidence of teosinte introgression into maize Is 
clearly seen in maize cobs throughout those parts of the 
valley where teosinte is most abundant. At harvest time 
the entire ear is brought to the granary, and for several 
weeks thereafter the ears are left to air dry in large piles 
around the courtyard. A sector of each of twenty piles 
representing twenty distinct fields of eight cultivators 
was studied. 'wenty-five cobs were separated from the 
pile and shelled. Approximately + of all the cob samples 
expressed the tripsacoid characteristic of a pronounced 
induration of the rachis and lower glume. All of the 
piles possessed tripsacoid cobs (Wilkes, 1968), although 


[ 300 | 


a few of the samples were free of tripsacoid cobs. 
Two of the fields where tripsacoid cobs had been har- 
vested were visited and teosinte was found to be abun- 
dant around the field margin. Although most of the 
teosinte plants had dispersed their seed and were dried 
and broken, several Fy hybrids were located which still 
possessed intact cobs. In one field, two teosinte backcross 
progeny of the F; maize Xteosinte hybrids were discov- 
ered, indicating that there issome reciprocal introgression. 
Despite the presence of reciprocal introgression, most 
of the gene flow is from teosinte to maize. Teosinte is 
a wild plant possessing the ability to disperse its seed as 
single rachis segments, while domesticated maize has a 
massive cob tightly enclosed by a husk system. Only 
those teosinte backcross segregates which possess the 
genetic control fora disarticulating rachis are able to dis- 
perse their seed. This factor is primarily responsible for 
the unidirectional flow of genetic material. The back- 
crosses to teosinte that segregate a more maize-like cob 
with non-disarticulating rachis disperse the entire spike 
as a whole. The numerous seed all germinating still at- 
tached to the cob are so crowded that they either choke 
each other out or develop into numerous spindly plants 
that fail to flower. In both cases, the genetic inheritance 
of the maize-like cob is lethal to a plant dependent on 
natural dispersal of its seed for survival. The selection 
tor the disarticulating teosinte pistillate spike and dis- 
tribution of single seeds protected by a rachis-segment, 
along with a large population of wild plants in the sur- 
rounding region, appears to act against the effects of 
maize introgression on the pistillate spike of teosinte. 


Study of maize ears 


The evidence for introgression of tripsacoid characters 
from teosinte is objectively measurable in the morpholo- 


[ 301 ] 


PLaTE LXNVI 


I \ | | 


Prare LXXVI. Evidence of teosinte introgression and 
maize <teosinte hybridization. A. The cob to the far 
left is a typical maize ear and shows evidence of intro- 
gression of tripsacoid segments associated with chromo- 
some 9, and possibly chromosome 4+. The cobs to the 
right are arranged in a series of increasingly pronounced 
tripsacoid characteristics. The extreme is the cob to the 
far right with its massive lower glumes and deep cu- 
pules. This cob, which was part of the maize harvest, 
is comparable in morphology to a maize backcross to the 
maize » teosinte hybrid. 7/10 actual size. B. This Fi 
maize < teosinte hybrid came from the same field as the 
cobsin A. Note the two-ranked rachis, the paired spike- 
lets, and the kernels forcing open the enclosing lower 
glumes; all typical characteristics of the Fi hybrid. 7/10 


actual size. 


[ 803 ] 


re 


ey of the maize cob (Galinat et al., 1956; Sehgal, 1963). 
The presence of a short, thick rachilla inclined to the 
axis, pronounced glume cushions, slightly upeurved 
glumes, comparatively soft rachis tissue, and well de- 
veloped central pith in the maize cobs of the fields sug- 
gests the introgression of segments of chromosome 9, 
and possibly chromosome 4, of teosinte. 

Might ears selected by the cultivator as seed ears from 
a field where maize X teosinte hybrids were known to be 
abundant were studied. These very productive ears were 
not highly tripsacoid, but they too showed evidence of 
teosinte introgression (rigid cob, straight rows, and in- 
durated glumes). Four of these ears were shelled and 
100 seeds from each ear were grown. Three cobs yielded 
all maize plants, but the fourth produced three maize X 
teosinte hybrids. This frequency of three maize X teosinte 
hybrids per 400 plants compares well with the number 
of maize X teosinte hybrids estimated to be present in the 
field and the abundance of highly tripsacoid cobs found 
in the total harvest. 

Maize Xteosinte hybrid seed on the predominantly 
maize-pollinated ear can not normally be distinguished 
morphologically from pure maize. Yet at Nobogame 
several ears from the same field were selected from the 
total pile because they possessed smaller than usual seeds. 
These small seeded ears uniformly yielded maize X teo- 
sinte hybrids when planted. It was found that if the ear 
is pollinated only by teosinte, the hybrid seeds are smaller 
than the few hybrid seeds found on a predominantly 
maize-pollinated ear. Thus there appears to be a chemi- 
cal feed-back mechanism (growth hormone/) between 
the developing seed and the cob. This postulated hor- 
mone might act to stimulate the conduction of food 
through the cob to the developing seed. 

Controlled pollinations of teosinte on the corn inbred 


[ 304 | 


A158 have produced seeds which are smaller and weigh 
less than self pollinated A158. This phenomena is not 
universal because controlled pollinations of teosinte on 
the New England Flint, Wilburs Flint, produced hybrid 
seed which are indistinguishable in size and weight from 
self-pollinated Wilburs Ilint. 


Discussion 

Maize and teosinte are fully fertile, and in the Nobo- 
game Valley the two hybridize naturally. Maize X teo- 
sinte hybrids are known from other regions in Mexico 
and Guatemala, but Nobogame is unique because hy- 
brids are in some cases cultivated because the inhabitants 
feel they improve the maize and in others the hybrids 
are unknowingly planted in the maize fields. 

Wild teosinte is abundant in the region, and hybrids 
are almost as abundant in the fields as on the margin of 
the fields. Because the method of cultivation of maize 
has not changed appreciably in the last hundred years 
this pattern of an abundance of teosinte hybrids on the 
seed ears Is suspected to have been widespread in the 
past in regions where it is almost extinct today. Such a 
pattern of hybridization with teosinte would account for 
the widespread tripsacoid characteristics now found in 
the races of maize on the Central Plateau (Wellhausen 
et al., 1950) and adjacent regions. This hybridization in 
the past would also explain why the teosinte of the 
Central Plateau is one of the most maize-like races of 
teosinte. 

The observations of the incorporation of teosinte seg- 
ments from chromosome 9, and possibly 4, are interesting 
also because Sehgal (1963) has shown that, under experi- 
mental conditions, incorporationof teosinte chromosomes 
9 and 4, either alone or in combination, in a uniform 
maize-inbred, increases the length of the cob. It is pos- 


— B05 | 


L >| 


Pirate LXXVII 


Evidence of tripsacoid characters in Nobogame cobs. A. These maize cobs are from a 
typical field where hybridization with teosinte was known to occur. Actual size. B. 
The short, thick rachilla inclined to the axis, the slightly upeurved glumes, and the 


pronounced glume cushions suggest the introgression of tripsacoid segments on chromo- 
some 9, and possibly chromosome 4, from teosinte. The rachis tissue is comparatively 
soft and the central pith is well developed. 3 actual size. 


tulated that, since increased cob length is selected for in 
seed ears, teosinte hybridization is of positive selection 
value in the maize of Nobogame. 


Summary 


1. 


A detailed analysis of the maize fields was made in 
the Nobogame Valley in southwestern Chihuahua, 
where the pattern of cultivation for maize has not 
changed appreciably in the last one hundred years. 
The maize grown around Nobogame is a five month 
type belonging to the race Cristalino de Chihuahua. 
The teosinte belongs to the race Nobogame and is 
abundant along the margin of maize fields and grow- 
ing wild in the non-cultivated parts of the valley. 


Maize and teosinte are not seasonally isolated as they 
are at most other sites where the two occur together. 
The mid-flowering times overlap in August and maize 
xXteosinte hybrids occur naturally. ‘Teosinte pollen 
is sufficiently abundant at flowering that many polli- 
nations on maize cobs are by a teosinte parent. Seed 
ears of maize have been shown to be contaminated by 
teosinte pollen resulting in the actual planting of 
maize X teosinte hybrids by the cultivator. 


The cobs of the maize planted in the valley show 
pronounced signs of tripsacoid germ plasm such as 
induration of the lower glume and a straight rigid 
cob. These characteristics are attributed to the intro- 
gression of tripsacoid genes via direct hybridization 
with teosinte foliowed by subsequent introgression 
from maize and teosinte backcross progeny. ‘The 
maize backcross progeny are usually harvested while 
the teosinte backcross progeny are usually left stand- 
ing in the field. Although introgression is reciprocal, 


[ 307 ] 


PLatTE LXNXVIILI 


Pirare LXXVIII. Teosinte-pollinated maize ears. 
A. When the maize ear is pollinated only by teo- 
sinte, the hybrid seed are smaller than the few 
hybrid seed found on a predominantly maize pol- 
linated ear, which are indistinguishable from the 
pure maize seed. The ear to the left is the inbred 
A158 and the ear to the right is a controlled polli- 
nation of teosinte pollen on the ear of A158. 7/10 
actual size. B. These ears are open pollinated ears 
from Nobogame. ‘The seed are of normal size on 
the ear to the left. The seed shelled from the ear 
to the right are smaller and all produced F); maize 
“teosinte hybrids. If the entire ear is pollinated 
by teosinte, none of the hybrid seed develop to the 
size of anormal maize kernel. This dwarfing effect 
only occurs if the entire ear is made up of hybrid 
kernels. Note also the tendency of the hybrid ker- 
nels to produce pointed seed, a character often 


found in pop corns. 7/10 actual size. 


[ 809 | 


the main flow of genes appears to be from teosinte 
to maize, since the genetic incorporation of a maize- 
like rachis results in the inability to disperse seed and 
the extinction of these maize introgressed teosinte 
plants. 


It is postulated that the hybridization of teosinte and 
maize on the Central Plateau in the past is compara- 
ble to the present hybridization of teosinte and maize 
at Nobogame. This hypothesis accounts for both the 
widespread presence of tripsacoid characteristics in 
the maize of the Central Plateau and the maizoid 
characteristics of the teosinte. The introgression of 
teosinte segments following maize X teosinte hybrid- 
ization would have been of positive selection value if 
it resulted in a larger ear. 


{ 310 | 


LITERATURE CITED 


Bukasov, S. 1926. Un hibrido de maize y Fuchlaena Mexicana. Fores- 
tal. (Mex.) 4: 38. 


Bukasov, S. 1930. Teosinte. in The cultivated plants of Mexico, 
Guatemala and Colombia. Bull. Appl. Bot. Genet. & Plant Breed- 
ing 47: 141-148. 


Collins, G. N. 1921. Teosinte in Mexico. Journ. Hered. 12: 339- 
350. 


Galinat, W.C., P.C. Mangelsdorf and L. Pierson. 1956, Estimates 
of teosinte introgression in archaeological maize. Bot. Mus. Leafl. 
Harvard Univ. 17: 101-124. 


Loépez y Parra, R. 1908. El teosinte. Mexico. 


Lumholtz, C. 1902. Unknown Mexico. Charles Scribner’s Sons, 
New York. 


Mangelsdorf, P.C. 1952. Hybridization in the evolution of maize. 
in J.W. Gowen (ed.), Heterosis, pp. 175-198. Iowa State College 
Press, Ames. 


Mangelsdorf, P.C., R.S. MacNeish, and W.C. Galinat. 1967. Pre- 
historic maize, teosinte and 7J'ripsacum from Tamaulipas, Mexico. 
Bot. Mus. Leafl. Harvard Univ. 22: 33-63. 


Mangelsdorf, P.C., and R.G. Reeves. 1959. The origin of corn III. 
Modern races, the product of teosinte introgression. Bot. Mus. 
Leafl. Harvard Univ. 18: 389-411. 


Sebgal, S.M. 1963. Effects of teosinte and ‘*Tripsacum’’ introgres- 
sion in maize, Bussey Institution, Harvard Univ. 


Wellhausen, E.J., L.M. Roberts and E. Hernandez-X, in collabora- 
tion with P.C. Mangelsdorf. 1952. Races of maize in Mexico. Bus- 
sey Institution, Harvard Univ. 


Wilkes, H.G. 1967. Teosinte: the closest relative of maize. Bussey 
Institution, Harvard Univ. 


Wilkes, H.G. 1968. Teosinte maize hybrids, Nobogame, Mexico. 
Maize Genetics Coop. Newsletter 42: 165-166. 


[ 311 | 


EARLY EIGHT-ROWED MAIZE FROM THE 
MIDDLE RIO GRANDE VALLEY, 
NEW MEXICO 


BY 
Watton C. Gatinat’, THEeoporE R. Remnuarr’, 
AND THEODORE R. FRISBIE* 


A MUTUAL interest of the maize phylogenist (senior 
author) and the archaeologist (junior authors) in the pre- 
historic spread of the eight-rowed race of maize, Maiz 
de Ocho, has made this and several other similar studies 
possible. Their respective interests, however, lie in dif- 
ferent aspects of the same problem. The evolutionary 
history of Maiz de Ocho involves the origin of the Corn 
Belt dent as well as many sweet corn varieties and its 
understanding could lead to the synthesis of even more 
productive strains in the future. It is an archaeological 
artifact of certain Indian cultures. The distribution of this 
race may also reveal some of the history of its possessors. 

In an earlier study it was found that the dates for this 


' Professor, Department of Environmental Sciences, University of 
Massachusetts, Waltham, Mass. and Honorary Research Associate in 
the Bussey Institution of Harvard University. 

? Asst. Prof., Department of Anthropology, College of William and 
Mary, Williamsburg, Va. 

> Ph.D. Candidate, Dept. of Anthropology, Southern Illinois Uni- 
versity, Carbondale, [I]. 

This investigation was supported in part by a grant (GB-15767) 
from the National Science Foundation to the senior author and in 
part by the Bussey Institution of Harvard University. 


[-313-} 


eight-rowed race generally became older toward the 
Southwest (Galinat and Gunnerson, 1968). A continuity 
in its distribution between the Plains and the Southwest 
was established by the discovery of sites for Maiz de 
Ocho in Chacuaco Canyon in southeastern Colorado 
(Galinat and Campbell, 1967). The date of A.D. 1140 
for this Colorado material leaves unsolved the problem 
on direction of spread because now the slightly older 
dates of A.D. 1040 have been discovered for this race at 
the Blain Site in Ohio (Galinat, 1969) and of A.D. 1125 
+70 at the Miller Site in Ontario (Kenyon, 1968). 

Documentation of the spread of Maiz de Ocho, like 
that of other cultural artifacts, depends upon the dis- 
covery of its oldest possible remains at certain critical 
locations. The present report on the oldest Maiz de Ocho 
yet discovered in North America has, therefore, particu- 
lar significance in revealing the spread of this race of 
maize. 

The radiocarbon dates herein reported were deter- 
mined gratuitously by Dr. Henry F. Nelson of the Re- 
search Laboratory of the Mobil Oil Corporation, Dallas, 
Texas. The date of 18188 B.C. (SM 1021) at the 
BR-45 Site came from an indirect association in debris 
between some wood and the cobs of Maiz de Ocho. The 
shallow and homogeneous nature of this debris seems to 
rule out any significant disparity between the age of the 
cobs and that of the analyzed wood. The more recent 
date of A.D. 870+168 (SM 1018) at Boca Negra Cave 
for this eight-rowed race came from wood which was 
charred in the same hearth with the cobs. At the third 
location for Maiz de Ocho discussed here, the Artificial 
Leg Sites, the dates of A.D. 550 to A.D. 700 for the 
cobs of this race were determined by combining data 
from ceramic analysis, palynology and archaeomagnetism 
(Frisbie, 1957). 


[ 314 | 


Tue LocaTIONs AND CULTURAL PHASES OF THE SITES 


THe BR-45 Srre 


The BR-45 Site is located on the west mesa at 6100 
feet about 124 miles west of the Rio Grande River at 
Albuquerque, New Mexico, while the Boca Negra Cave 
and Artificial Leg Sites are elevated at about 5100 feet 
on the first terrace closer to the river. The other maize- 
containing sites previously reported in central New 
Mexico are on the same western side of the river and to 
the southwest of the Middle Rio Grande Valley (Plate 
LXXIX). 

The west mesa is now dissected by arroyos sloping in 
a west-to-east direction toward the river. The arroyos 
cut through dune-covered ridges which were the occupa- 
tion sites. During a number of wetter years than at 
present, maize was probably grown on the flood plains 
where these more modern arroyos now occur. 

The BR-45 Site occupies the south slope of one of the 
sandy ridges just below its crest. It consists of two small 
pithouses with outside living areas to the east of the 
dwellings, as described in detail by Reinhart (1967a). 

The cultural affinities at the BR-45 Site were of the 
Alameda Phase which is an early Basketmaker [11 mani- 
festation. Its identification is based on typological com- 
parisons, including ceramics, to known Basketmaker II] 
material. These identifications (Reinhart, 19672) suggest 
that the dunes were stabilized approximately 2000 to 
1500 years ago during a period of increased moisture. 
This dating also appears to fit the moist period postulated 
as being between the Fairbank and Whitewater droughts 
(Antevs, 1955) as well as the radiocarbon date of 18+ 
138 B.C. for the BR-45 maize, as will be described later. 


PLATE LXXINX 


| ] | | 


Cebollita Cave 


I 
1 
! 
1 
i 
| 
| 
| 
| 
| 
] 
1 
1 
I 
] 
| 
1 
I 
! 
1 
I 
| Bat Cavee 


a 
Tularosa Cave 


| 
| 
! 4 
= 
a Sind Boca Negra. 
Cave 
4 
ae ee zt 
oo : 
f I Ns N 
oO 50 mi. 
a _ 4 
S NO vA C i, 
/ 
| | | | | | | 
109 108 107 106 105 104 103 


The enlarged insert shows the three site areas, BR-45, Boca Negra Cave, and 
Artificial Leg, which are studied in this paper. Other maize-containing sites in 
New Mexico are indicated to the southwest of this Albuquerque area. 


[ 316 | 


Boca Necra Cave 


Boca Negra Cave is situated on the northeast side of 
a voleanic cone near the city of Albuquerque. The cave 
measured 274 feet deep, 205 feet wide and 6 feet high 
before excavation. The details of the excavations are 
described by Reinhart (1968). 

Because the terrain around the cave and for a mile to 
the river is rocky and otherwise unsuited for agriculture, 
the site probably never had a permanent agricultural 
settlement. Nevertheless along record of maize and other 
artifacts was left apparently by migrants who camped at 
the cave for a short time and then moved to other Bas- 
ketmaker III agricultural sites along the river such as 
those at Artificial Leg. 

The stratigraphy at Boca Negra Cave has a bearing 
on our understanding of the evolution of southwestern 
maize. Its succession includes material which is both 
older at the lower level and more recent at the upper 
level than that found at the BR-45 Site. The earliest 
culture present was of the late Atrisco Phase (Campbell 
and Ellis, 1952) which dates prior to 1000 B.C. and is a 
local Cochise manifestation. While incipient agriculture 
may have been practiced, there is no direct evidence of 
it either at Boca Negra Cave or at the west mesa sites 
at this early time. The Atrisco Phase is preceramic and 
is represented here only by lithic tools. While the next 
Phase, Rio Rancho, was also preceramie, it contained 
cobs of Chapalote, the indigenous race of the Southwest 
and Mexico which will be described later. Both the ce- 
ramics and a new race of maize, Maiz de Ocho, appear 
in the Alameda Phase, an early Basketmaker III culture 
dated at A.D. 370+168 at the Boca Negra Cave Site. 
This stratigraphy extending over a 8000 year period to- 
gether with the racial identity of the maize present are 
given in Table IIT. 

[ 317 | 


THe ARTIFICIAL LEG SITES 


The three excavated Artificial Leg Sites are also lo- 
cated near the river about 12 miles north of Boca Negra 
Cave and just to the west of Sandoval (Corrales), New 
Mexico (Frisbie, 1967). The initial settlement of Site I 
occurred during the same drought period around A.D. 
550 when Maiz de Ocho was introduced into the already 
established culture of Boca Negra Cave. The settlers at 
Artificial Leg also appear to have migrated from their 
original home on the mesa (BR-45) to these sites near 
the river in order to have moist soil necessary for the 
growth of their maize. 

The shelters at these Middle Rio Grande Valley sites 
were pithouses. Their structure coincides more closely 
with that of the Mogollon custom in the South than with 
that of the Anasazi of the North. They were nearly cir- 
cular in outline with either a flattened or concave east 
side. The roofs were usually supported by four posts. 
Floor features consisted of a centrally located, circular, 
collared hearth, deflectors, ash pit, ladder holes, and an 
easterly oriented ventilator shaft. Other floor features 
included depressions and storage pits, many widened at 
their bases. 

At Artificial Leg, each site consisted of a group of at 
least four such pithouses. One of these structures was 
larger than the others, being 26 feet in diameter rather 
than the approximately 16 feet commonly found. This 
larger pithouse, more properly termed ‘‘kiva,’” served 
ceremonial purposes, as indicated by its s¢tpapu or ‘‘open- 
ing to the underworld,” by its foot drums and by its 
location apart from the other pithouses. The develop- 
ment of these ceremonial structures again reflects Mo- 
gollon customs. 

The appearance of the new productive races of maize, 


[ 318 | 


Maiz de Ocho and Pima Papago, with the Mogollon 
type of pithouse, especially the ceremonial type, indi- 
cates that the two may have diffused together from the 
South. The short tenure of these races on the mesa which 
was terminated by drought may not have allowed time 
for the elaboration of appropriate ceremonialism. 

The evidence from ceramics at these sites near the 
river indicates contacts with both the southern and 
northern cultures. Potsherds from the Alma Plain which 
originated with the Mogollons from the South were 
found with mineral and carbon painted affiliates of Lino 
Gray representing Anasazi wares from the North. While 
the data on ceramics do not in themselves verify the 
route of Maiz de Ocho, they do reveal that trade was 
well established between the South and North. This new 
race of maize would probably have been included in some 
of the transactions. 


THE MaIZE 


Chapalote is the indigenous race of maize of the area. 
It traces back to preceramic times during the Rio Rancho 
phase at Boca Negra Cave. About 125 miles to the south 
at Bat Cave, New Mexico, Chapalote was dated at 2300 
B.C. where its evolution eventually involved a second 
element, the wild relative teosinte, at 500 B.C. (Mangels- 
dorf and Smith, 1949; Mangelsdorf et al., 1967). 

Chapalote may be identified either by its kernels or by 
its cobs. Its kernels are deeper than wide in shape and 
generally small and flinty although a floury form was 
selected in Basketmaker times. Its cupules (depressions 
in the cob axis (rachis) immediately above the attach- 
ment point of each pair of pistillate spikelets) are narrow 
(3 to 4 mm. wide) and the width and height (internode 
length) are usually of about equal dimensions. Evolved 
Chapalote has 12 to 16 rows of kernels while the earliest 


[ 319 | 


TABLE 1. 


MAIZE FROM THE BR-45 SITE, NEW MEXICO 
No. Kernel] w=Cupule | €Internode | Cupule 
Sample Number Rows Width(mm) | Length(mm) | Shape w/2 | Race 
BR-45/M1-1 12 5.6 3.5 1.6 Pima Papago 
. -2 12 a6 pee tee : " 
-3 12 5.4 3.5 leo . " 
’ -4 12 5.6 336 1.6 ’ . 
BR-45/M2 8* 6.0 3.0 2.0 Mafz de Ocho 


*Row number estimated on the basis of cob fragments consisting of at least two parellel rows 


of cupules. 


w/L= cupule width/cupule internode length. 


[ 820 ] 


Chapalote, such as that from Bat Cave, has eight kernel 
rows. The race Reventador is very similar to evolved 
Chapalote except for having colorless instead of brown 
pericarp. Because no distinction between Reventador 
and Chapalote was possible in this archaeological ma- 
terial, the name of the older race, Chapalote, is used. 

The race of particular concern, Maiz de Ocho, was the 
third major element involved in the evolution of maize 
in the Southwest. Previously this race was thought to 
have arrived in New Mexico at about A. 1). 700, the date 
of its earliest remains at Tularosa Cave (Cutler, 1952). 
Apparently it appeared at about the same time 90 miles 
northward at Cebollita Cave (Plate 23 F of Galinat and 
Ruppé, 1961). The earlier dates of 18+138 B.C. for 
Maiz de Ocho 75 miles to the northeast at BR-45 and 
A.D. 370168 at Boca Negra Cave do not negate the 
overall evidence for a southern origin. The next oldest 
date for this race is A.D. 200-850 previously reported 
for a site in 'amaulipas, Mexico (Mangelsdorf et al., 
1967). The Tamaulipas material may be on a parallel 
pathway leading to the Davis Site in Texas which had 
this eight-rowed race at A.D. 800-1000 (Jones, 1949). 
The Maiz de Ocho ancestral to that from the Albuquer- 
que area of New Mexico more probably came directly 
through Chihuahua and Sonora, as indicated by four 
caves in these Mexican states yielding prehistoric re- 
mains of this race (Mangelsdorf and Lister, 1956). Appar- 
ently southern United States had multiple connections 
through the Sierra Madre Occidental of northern Mexico 
from some common source, perhaps on the west coast of 
southern Mexico. Maiz de Ocho is thought to have been 
derived originally from Colombia, South America (Gali- 
nat and Gunnerson, 1963). 

The kernels and cob of Maiz de Ocho are distinctive 
from those of Chapalote. They occur in only eight rows 


[ 321 ] 


(borne in pairs at four ranks of cupules) instead of the 12 
to 16 rows which characterize the older race. The shape 
of kernel is also different, being wider than deep rather 
than deeper than wide. In addition, the shape of the 
cupule is definitive. The ratio of cupule width/internode 
length is about two in Maiz de Ocho in contrast to the 
more narrow cupules of Chapalote, especially near the 
tip of the cob where the ratio is close to one. 


THe MaIze FROM THE BR-45 SIre 


The maize from BR-45 consisted of five broken and 
eroded cobs. ‘The kernel row number, cupule width and 
internode length of these specimens was measured in 
order to establish their racial identity (Table I). One of 
the cobs represented the race Maiz de Ocho and the other 
four were from its hybrid, Pima Papago. The eight- 
rowed cob could be a segregate out of the hybrid. In 
any case the hybrid was most abundant. The early ac- 
ceptance of the hybrid may be due to its rapid adaptation 
through the segregation of favorable genes from its in- 
digenous parent, Chapalote. he increased vigor and 
productivity associated with such a wide cross must have 
promoted its spread, probably far ahead of the Maiz de 
Ocho parent. 

The cultivation of the productive races of maize, Maiz 
de Ocho and Pima Papago by the people at BR-45 
seems to have bound their way of life to farming. Serv- 
ing these people as a major source of food, they became 
dependent upon it. Subsequently after a protracted dry 
period they abandoned the mesa for sites nearer the river 
which provided the moist soil necessary for the growth 
of their maize. After abandonment, the cobs of Maiz de 
Ocho and Pima Papago and other debris on the ridge 
sites became buried under the sands of time until the 
present excavation. Previously when their ancestors with 


[ 822 ] 


MAIZE FROM BOCA NEGRA CAVE, NEW MEXICO 


TABLE II. 


No. Kernel] Cupule Internode Cupule 

Sample Rows Width(mm)| —Length(mm) | Shape w/ Race 
BB-1/MI1-1 12 3.5 3.5 1.0 Chapalote 

nH 12 4.0 2.8 1.4 Chapalote Mix 

"23 12 5.0 $2 1.6 Chapalote Mix 

n 4 12 3.5 3.5 1.0 Chapalote 
BB-1/M2 10 6.5 3.0 22 Matz de Ocho 
BB-1/M3 12 4.6 3.0 1.5 Mafz de Ocho Mix 
BB-1/M4-1 10 6.5 a7 1.8 Matz de Ocho 

. -2 14 2.5 358 1.8 Pima Papago 
BB-1/M5-1  c. 10 5.0 a3 1.4 Maiz de Ocho Mix 

m 22 6. 10 5.5 3.5 1.6 Mafzude Ocho Mix 
BB-1/M6-1 cc. 10* 5.7 4.2 1.4 Matz de Ocho Mix 

"22 OG, 8 5.4 4.0 1.1 Matz de Ocho Mix 

a rn 8 5.6 4.1 1.4 Mafz de Ocho Mix 

a 8 5.5 4.1 Le Mafz de Ocho Mix 
BB-1/M7-1 12 4.6 4.0 12 Chapalote Mix 

me =? 12 3.1 3.1 1.0 Chapalote 

o. 73 12 3.2 3.2 1.0 Chapalote 
BB-1/M8-1  c. 12 3.5 3.5 1.0 Chapalote 

<= 12 3.4 a5 0.97 Chapalote 

" 12 231 22 0.94 Chapalote 
BB-1/M9-1 c. 10* 6.5 2.6 2.5 Matz de Ocho 

age 12* 7.3 3.0 24 Pima Papago 

"23 10* 6.0 a3 1.8 Mafz de Ocho 

"  =4 12% 5.3 3.6 1.5 Pima Papago 
BB-1/M10-1. c. 10 6.7 4.2 1.6 Matz de Ocho Mix 

" -2 ¢. 6 8.1 4.6 1.8 Mafz de Ocho Mix 
BB-1/M11 14 4.5 4.5 1.0 Chapalote 


*Row number estimated on a basis of fragments consisting of at least two parallel rows of cupules. 


c. - carbonized cob. 


[ 328 ] 


less productive food plants encountered similar drought 
conditions on the mesa, they merely adapted by shifting 
their subsistence patterns to include more wild food 
rather than seek a more suitable farming area. 

The early radiocarbon date (18+188 B.C.) and the 
major cultural traits at BR-45 indicate that the races of 
maize, Maiz de Ocho and Pima Papago, located there 
were originally derived from the southern Mogollon cul- 
ture rather than from the Anasazi of the San Juan drain- 
age area to the North. That is, the age of the Basket- 
maker cultural level at BR-45 and the other nearby sites 
discussed in this paper is intermediate between that of 
the older Mogollon cultures of the South and the more 
recent San Juan cultures of the North. This suggests 
that the Middle Rio Grande Valley was an area of tran- 
sition from South to North. 


THe Maize rrom Boca NeGra Cave 


Before the introduction of Maiz de Ocho, the indige- 
nous race of maize, Chapalote, occurred in Basketmaker 
II (preceramic) times about twelve miles away toward 
the river at Boca Negra Cave. Later the people presuma- 
bly moving from the drying mesa area (BR-45) left their 
Maiz de Ocho and Pima Papago maize during the early 
Basketmaker II1 period (the end of the Alameda Phase). 
The radiocarbon date of A.D. 370+168 for this new 
maize at the cave is a jump of over 400 years over that 
reported for its collection at BR-45, only 124 miles away 
but 1000 feet higher on the west mesa. 

There were 24 eroded cobs, eight of which had been 
carbonized, from Boca Negra Cave. The excavation was 
in stratigraphic layers from the surface to a depth of 40 
inches. ‘he cobs represent two distinct races, Chapalote 
and Maiz de Ocho and their hybrid, Pima Papago, as 
well as some intermediate types classified as ‘‘mixed”’ 


[ 324 | 


TABLE Ill 


THE STRATIGRAPHY AND MAIZE FROM BOCA NEGRA_ CAVE 


: Race of Maize 
Maize 
Phase of Sample : 
x1 0 
Cultural Association Number OL mre 
BB-1 2 =| 81 6] O 
(e) E ro] Oo 0 
ire; . [a (by bs © | 
5 o e ~ se 
O Ol =] =] s 
Pueblo IV to Historic M1 2 2 010} 0 
Pueblo IV M2 0 0 Oo {O07}; 1 
Pueblo III to IV M3 0 0 O11] 0 
Pueblo II to IV M4 0 0 170} 1 
1 
Early Alameda M9 0 0 21/0]; 2 
Rio Rancho to Alameda M5 0 0 0 |21| 0 
"W " " " M6 0 0 0 4 0 
Rio Rancho to Early Alameda M8 3 0 0 |;0] 0 
Rio Rancho“ mz {2 |1 | 0 ]o]o 
Atrisco™ : 0 |o |oflojo 
1 - Early Basketmaker II], dated A.D. 370+ 168 
2 - Basketmaker II, preceramic 
3 - Local Cochise, no direct evidence for agriculture prior to 1000 B.C. 


Two additional maize samples (BB-1/ M10 and BB-1/M11) of unknown 


cultural association are not included. 


[ 825 ] 


which probably represent the backcross or segregating 
progenies derived from the hybrid. The typical hybrid 
type (Pima Papago) combined the higher kernel row 
number (12 rows) of Chapalote with the wider cupules 
(6 to 10 mm.) of Maiz de Ocho. When the mixed modi- 
fication is toward Maiz de Ocho, the specimen is scored 
as Maiz de Ocho Mix. When the direction is that of 
Chapalote, it is identified as Chapalote Mix. The data 
upon which the races and forms were identified and the 
phase of cultural association for the various specimens 
are given in Tables [I and III. 


THE MaIZE FROM THE ARTIFICIAL LEG SITES 


All of the 13 cob fragments of maize from the Artifi- 
cial Leg Sites were completely carbonized. ‘Ten of these 
came from Site I dated at A.D. 550 to A.D. 700, two 
from Site II dated at A.D. 750 to A.D. 800, and one 
from Site III dated at A.D. 870 to A.D. 980. The hy- 
brid race, Pima Papago, predominates and the larger 
collection (Site I) contains a few specimens similar to its 
parents, Chapalote and Maiz de Ocho. The latter appear 
to be parental-type segregates from the hybrid rather 
than independently derived pure entities (Table IV). 

Pima Papago is presumed to have been more success- 
ful at first than its introduced parent, Maiz de Ocho and, 
thereby, to have traveled northward from the Sierra 
Madre Occidental at a faster pace. It is unfortunate that 
more specimens were not available from Site III. The 
single specimen available was of a modified form of Maiz 
de Ocho which may well represent a segregate from the 
hybrid. 

Apparently in the more northerly climates natural 
selection favors the eight-rowed derivatives. Further 
north across the boundary into southeastern Colorado, 
the Chapalote type segregates almost disappear while 


[ 826 | 


TABLE IV 


MAIZE FROM THREE SITES AT ARTIFICAL LEG, NEW MEXICO 
No. Kernel | Cupule Internode Cupule ~ 
Sample Number Rows Width(mm) | Length(mm)| Shape w/ Race 
Site 1, Pithouse 6, West Cist Floor: A.D. 550 to A.D. 700 
BAL-35/1 14 6.9 oi) 2.30 Pima Papago 
 2a/2 12 ao Ale 1.96 Pima Papago 
" 35/3 12 4.5 Sie 1.43 Chapalote 
" 35/4 8 10.0 3.2 3.10 Mafz de Ocho 
W 35/5 12 51 3.5 1.46 Pima Papago 
"35/6 14 4.8 3.4 1.4] Chapalote Mix 
H 95/7 10 5.2 3.6 1.45 Pima Papago 
95/8 14 3.8 2:0 1.90 Pima Papago 
e O57 16 5.9 3.0 1.97 Pima Papago 
" 35/10 ]2 4.9 3.1 1.58 Chapalote 
Site Il, Pithouse 4, Fill: A.D. 750 to A.D. 800 
BAL-35/11 12 5.5 | 3.5 1.57 Pima Papago 
"25/12 14 5.7 3.7 1.54 Pima Papago 
Site Ill, Pithouse 1, Fill: A.D. 870to A.D. 930 
BAL-35/13 10 6.1 | 2.7 2.26 Maiz de Ocho Mix 
[ 827 ] 


Maiz de Ocho germplasm predominates. Here in the 
Chacuaco Canyon area, Chapalote was reduced to only 
two percent while Maiz de Ocho occurred at 22.8 per- 
cent, Maiz de Ocho Mix at 36.9 percent and the inter- 
mediate hybrid type (Pima Papago) at 386.3 percent from 
a total of 201 cobs (Galinat and Campbell, 1967). As 
the distribution progressed onto the Plains and north- 
eastward, the frequency of pure Maiz de Ocho increased. 
Nevertheless, even the northern flints may exhibit some 
degree of Chapalote germplasm derived from their an- 
cestral mixing in the Southwest such as is manifest by 
the flinty kernels or the occasional kernel row numbers 
of over eight. 


SUMMARY 


The radiocarbon date of 18188 B.C. is the earliest 
North American date yet reported for the introduced 
race of maize, Maiz de Ocho, or its hybrid, Pima Papago, 
involving the indigenous race, Chapalote. The early date 
comes from the BR-45 Site about 124 miles to the west 
and 1000 feet higher than the Albuquerque area of the 
Middle Rio Grande Valley, New Mexico. Older remains 
of the original pure form of Maiz de Ocho probably will 
be discovered in Chihuahua and Sonora, perhaps near 
the valleys draining to the west coast of Mexico. From 
there, it is presumed to trace to South America where 
it has botanical affinities with the Colombian race, Ca- 
buya (Roberts et al., 1957). The early presence of the 
hybrid Pima Papago in New Mexico rather than the 
pure Maiz de Ocho parent can be attributed to an in- 
creased adaptability, vigor and productivity associated 
with outcrossing to the indigenous race, Chapalote. The 
eight-rowed parent remains in a semi-hybrid condition 
in the form of Harinoso de Ocho in northwestern Mexico 
(Wellhausen et al., 1952), while it has assorted out to an 


| $28 | 


increasing pure race as it spread onto the Plains and to 
the Northeast in its well known forms of northern flour 
and northern flint. ‘The presumed Colombian ancestor, 
Cabuya, is a highland race, and this might account for 
some pre-adaptation to high altitudes and northern lati- 
tudes in United States, as exemplified by the early distri- 
bution of Maiz de Ocho (Galinat and Gunnerson, 1968). 
The Maiz de Ocho from the BR-45 Site was moved 
by its cultivators to sites with moist soil along the river 
after a protracted dry period made the mesa unsuitable 
for the growth of their maize. At the lower elevations, 
it became further mixed with the older indigenous race, 
Chapalote, in Basketmaker II] times at the end of the 
Alameda Phase. The radiocarbon date of A.D. 870+ 
168 marks the introduction of this eight-rowed maize 
into the strata at Boca Negra Cave. A somewhat later 
date of A.D. 550 to A.D. 700 follows for this race just 
north of Albuquerque at the Artificial Leg Sites. By 
A.D. 1040 Maiz de Ocho had reached the Blain Site in 
Ohio (Galinat, 1969) and by A.D. 1125+70 the Miller 
Site in Ontario (Kenyon, 1968). While the frequency 
and purity of Maiz de Ocho increased during its spread to 
the Northeast, some traits of Chapalote derived from an 
ancestral mixing inthe Southwest may still be recognized 
in the Canadian and New England flint corn varieties. 


[329 ] 


LITERATURE CITED 


Antevs, E. 1955. Geologic-climatic dating in the West. American 
Antiquity 20(4): 317-335. 


Campbell, J.M. and F.H. Ellis. 1952. The Atrisco Sites: Cochise 
manifestations in the Middle Rio Grande Valley. American An- 
tiquity 17(3): 211-221. 


Cutler, H.C. 1952. A preliminary survey of plant remains of Tula- 
rosa Cave. Chicago Museum of Natural History. Fieldiana: Anthro- 
pology 40: 461-479. 


Frisbie, T.R. 1967. The excavation and interpretation of the Artifi- 
cial Leg Basketmaker I1I—Pueblo I Sites near Corrales, New Mex- 
ico (unpubl. M.A. Thesis, Univ. of New Mexico, Dept. of Anthro- 
pology). 


Galinat, W.C. 1970. Maize from the Blain Site in Ohio: in O.H. 
Prufer. Archaeology of the Blain Site. Monograph, Western Re- 
serve Univ. Press. (In press. ) 


—— and R.J. Ruppé. 1961. Further archaeological evidence on the 
effects of teosinte introgression in the evolution of modern maize. 
Bot. Mus. Leafl. Harvard Univ. 19: 163-181. 


~and J.H. Gunnerson. 1963. Spread of eight-rowed maize from 
the prehistoric Southwest. Bot. Mus. Leafl. Harvard Univ. 20: 
117-160. 


—— and R.G. Campbell. 1967. The diffusion of eight-rowed maize 
from the Southwest to the Central Plains. Mass. Agric. Exp. Sta. 
Mem. Ser. 1: 1-16. 


Jones, V.H. 1949. Maize from the Davis Site; its nature and inter- 
pretation: in H.P. Newell and A.D. Krieger, The George C. Davis 
Site, Cherokee County, Texas. Soc. for Amer. Archaeology Memoir 
5: 239-249, 


Kenyon, W.A. 1968. The Miller Site. Occasional Paper 14, Royal 
Ontario Museum, Toronto. 


Mangelsdorf, P.C. and C.K. Smith, Jr. 1949. New archaeological 
evidence on evolution in maize. Bot. Mus. Leafl. Harvard Univ. 
13: 2138-247. 


[ 880 | 


Mangelsdorf, P.C. and R.H. Lister. 1956. Archaeological evidence 
on the evolution of maize in northwestern Mexico. Bot. Mus. Leafl. 
Harvard Univ. 17: 151-178. 


—— , R.S. MacNeish and W.C. Galinat. 1967. Prehistoric maize, 
teosinte, and Tripsacum from ‘Tamaulipas, Mexico. Bot. Mus. Leafl. 
Harvard Univ. 22: 33-63. 


Reinhart, T.R. 1967a. The Alameda Phase: An early Basketmaker 
III culture in the Middle Rio Grande Valley, New Mexico. South- 
western Lore 33: 24-32. 


——, 1967b. The Rio Rancho Phase: A preliminary report on early 
Basketmaker culture in the Middle Rio Grande Valley, New Mex- 
ico. American Antiquity 32: 458-470. 


——., 1968. Late Archaic Cultures of the Middle Rio Grande Valley, 
New Mexico. Unpublished doctoral dissertation, University of New 
Mexico, Albuquerque. 


Roberts, L.M., U.J. Grant, R. Ramirez E., W.H. Hatheway and 
D.L. Smith in collaboration wilh P.C. Mangelsdorf, 1957. Races of 
Maize in Colombia. Nat. Acad. Sci.-Nat. Res. Council Pub. 510, 
133° Pps 


Wellhausen, E.J., L.M. Roberts and E. Hernandez X, in collabora- 
tion with P.C. Mangelsdorf. 1952. Races of Maize in Mexico. The 
Bussey Institution of Harvard Univ., Cambridge, Mass. 223 pp. 


[ 381 ] 


334 


BOTANICAL MUSEUM LEAFLETS 
HARVARD UNIVERSITY 


CambBrincr, Massacuuserrs, June 29, 1970 VoL. 22, No. 


ETHNOGYNECOLOGICAL NOTES IN THE 
HARVARD UNIVERSITY HERBARIA 


BY 
Srri von Reis ALTSCHUL 


PLANts used by primitive societies for the maintenance 
or restoration of health have drawn the attention of 
medical researchers in modern times. Those species 
which, however vaguely, have offered hope for relief 
from cancer, heart disease, mental illness and diverse 
metabolic disorders have aroused probably greatest in- 
terest up to the present. Among the plants which remain 
for the most part unexplored as to their pharmacological 
potentials are species which have been employed in con- 
nection with the functions and diseases of the reproduc- 
tive tract of the human female. Asa branch of ethno- 
medicine, this field might be called ethnogynecology. 
I use the term here to include, as well, ethnoobstetrics, 
embracing the practices surrounding pregnancy, labor 
and the puerperium in primitive cultures. 

A number of species of plants previously little known 
for their involvement in ethnogynecology recently has 
been brought to light as the result of a large-scale search 
carried out during the last decade at Harvard University. 
This paper is one among several which have dealt with 
medicinal folklore in general,’ with psychopharmacolo- 
ey,’ with unusual food plants’ and with plants used to 
treat children’s diseases,* all based on data retrieved 


from the survey. 
[ 383 ] 


In 1962, Dr. Richard Evans Schultes and I set upa 
search for botanical folklore of potential interest to health 
and medical sciences.* Projects with this objective are 
not new. Ethnobotanical information has been gathered 
traditionally from both old and new writings in anthro- 
pology and botany, from archaeological materials and 
from field work. We believe that our project was unique 
in that it relied exclusively on mainly unpublished data 
from labels accompanying herbarium specimens. In five 
years, a sheet by sheet examination of over 2,500,000 
specimens of flowering plants in the combined herbaria 
of the Arnold Arboretum and Gray Herbarium of Har- 
vard University was completed. ‘These specimens span a 
century and a half and are worldwide in representation. 
Irom these repositories the harvest amounted to nearly 
7,000 notes from over 5,000 species. Materials ranging 
from magic to chemistry were recorded there, wherever 
was some hint of the presence of biodynamic agents. All 
species were checked in Uphof’s Dictionary of Hconomic 
Plants’ in order to eliminate from our compilation al- 
ready well known uses. One exception which was made 
to this procedure was in conserving any note which ex- 
tended the geographic representation of a known use. 

The search was undertaken in the belief that the major 
herbaria of the world represent untapped reservoirs of 
vital data. In some instances, they might provide the 
only remaining clues to the materia medica and nutritional 

* The project was supported in turn by Smith, Kline and French 
Laboratories, the National Institute of Mental Health, and the Eli 
Lilly Research Laboratories. It was sponsored by the Botanical Mu- 
seum of Harvard University. The author is indebted to Dr. Schultes 
for his encouragement and guidance. She is grateful also to Profes- 
sors Richard A. Howard and Reed C., Rollins, Directors respectively 
of the Arnold Arboretum and Gray Herbarium of Harvard University, 
for their generous permission to use these two herbaria. This paper 


was presented on August 31, 1969, at the XI International Botanical 
Congress (under Ethnobotany), in Seattle, Washington. 


[ 384 ] 


patterns of peoples now extinct or whose cultures have 
been absorbed or destroyed by civilization. In other in- 
stances, they might serve to point out species which have 
not been used by man but which, on the basis of field 
observations, lead one to believe that they could be de- 
veloped as economic plants. It was felt that the accelerat- 
ing expansion of urbanization and forced constriction of 
natural areas make it urgent to gather and study all 
possibly useful species before living representatives in 
nature inadvertently are extinguished. 

Out of the entire body of notes extracted from the 
herbarium search, about 100 notes fall within the class 
of ethnogynecology as defined above. These 100 or so 
notes represent as many field collections and nearly as 
many species. It is likely that many additional species 
from our notes are employed for the same purposes but 
are unrecognizable as such on the basis of the field notes 
alone. For example, in the newly completed index to 
our notes, there are over 400 species described only as 
‘‘medicinal plants’’. There are almost 200 species which 
are reputed analgesics. Some 85 species are used for 
stomach disorders. Increasingly fewer numbers are em- 
ployed as astringents; against abdominal spasms; as 
aphrodisiacs; for treating anemia; in baths; and as 
anti-convulsants. 

Of the roughly 100 notes which are the concern of this 
paper, 389 have to do with the period after labor known 
as the puerperium. ‘Twenty-one per cent relate to treat- 
ment of venereal disease, not always indicating, it must 
be acknowledged, whether for males or females. Eleven 
per cent deal with menstruation. Nine per cent refer to 
‘‘female disorders’’. Eight per cent have to do with 
labor or childbirth. Another 8% relate directly to birth 
control. The remainder concerns galactagogues and 
plants used during the course of pregnancy. Needless 


to say, some notes overlap more than one category. 

Fifty families of flowering plants are represented 
among these notes. Thirty of these families have yielded 
one note each, which suggests a scattered distribution 
of ethnogynecological uses among the flowering plants. 
‘Twelve families produced two notes each. The Borag- 
inaceae and Labiatae are represented each by three notes, 
the Gramineae by four, the Liliaceae five notes. The 
Leguminosae had six, Huphorbiaceae eight, Compositae 
12 and Rubiaceae 18. 

At least 35% of the plants are said to be eaten or, in 
some form, drunk. Sixteen per cent are made into de- 
coctions or infusions, most of which probably are also 
orally administered. Fourteen per cent are employed 
externally, including in baths. Two per cent are inserted 
vaginally. Thirty-three per cent of the notes do not re- 
veal the modes of employment. 

Seventy per cent of the ethnogynecological notes indi- 
cate the parts of plants used. Roots are said to be used 
in nearly half of these notes. Leaves are designated in 
almost a third. The rest of the notes cite, in order of 
decreasing importance, barks, the entire plant, fruits, 
flowers and latex. 

Of the 87 genera represented among these ethnogyn- 
ecological notes, 33 are virtually unknown economically. 
Three quarters of the species are similarly unknown for 
economic purposes of any sort. 

I should like to cite some of the field notes associated 
with species of economically obscure genera. As already 
stated, the largest group of ethnogynecological notes 
concerns the puerperium. In this category, sundry spe- 
cies can be cited from the Philippine Islands. For ex- 
ample, Schizostachyum Lumampao (C. O. Frake 567, 
Gramineae) and Nadsura scandens (C. O. Frake 578, 
Schisandraccae) are used as postpartem medicines. The 


[ 886 ] 


roots of Anema glomerata (C. O. Frake 569, Myrwstica- 
ceae) and Phytocrene Blancoi (C. O. Frake 502, Icacina- 
ccae) are boiled and drunk during puerperium. ‘The roots 
of Melicope monophylla (C. O. Frake 674, Rutaceae) are 
pounded and drunk, while those of Aphana sp. (C.O. 
Frake 38140, Ieacinaceae) are applied in some manner. 
The bark of Neonauclea formicaria (C. O. Frake 566, 
Rubiaceae) is boiled and drunk. The genus Cyrtandra 
(C. Cumingti, R. B. Fox 5052; Cyrtandra sp., M.D. 
Sulit 9974, Gesneriaceae) is used in association with 
bleeding following childbirth. A noteworthy claim is 
made for an indeterminate species of Polyosma (M. D. 
Sulit 3443, Saxifragaceae): ‘‘Decoction of roots given 
to women who gave birth—after delivery women can 
immediately walk—according to native’’. 

From South America, one might cite Tournefortia 
brevilobata(W. H. Camp 2560, Boraginaceae): in Kcua- 
dor, ‘‘Infusions of lvs. drunk by women a day or two 
after childbirth to ‘clean everything out’ ”’. 

In association with the puerperium, one could include 
reputed galactagogues, such as Mpigynum Maingayt 
(Ismail & Millard s.n.: K. L. No. 187, Apocynaceae) 
from Malaya. From Indochina, the following was re- 
ported tor Desmos Hancei (Poilane 1184, Annonaceae): 
‘«. les Indigénes...emploient les rameaux et feuilles 
pour faire un breuvage qu’ils font prendre au femme 
aprés accouchement pour augmenter la secrétion lactée, 
ils donnerait également des fruits comestibles”’. 

The second largest group of ethnogynecological notes 
gathered relates to venereal disease. At least two species 
are represented from the Solomon Islands: /aradaya sp. 
GS. FF. NKajewshi 2543, Verbenaceae) ‘‘... in common 
with other vines is used for gonorrhea, the bark is macer- 
ated with water, the resulting concoction being drunk’. 
Cremastus sceptrum (Williams & Assis 7543, Bignonia- 


[ 837 | 


ceae) is supposed to be the source of a root-derived syrup 
‘*Good for syphilis”’. 

There are various examples from the New World: in 
KE] Salvador, Campelia zanonia (P. C. Standley 19320, 
Commelinaceae) is a *‘Remedy for gonorrhoea’. In Bo- 
livia, Pothomorphe peltata (J. Steinbach 5536, Pipera- 
ceae) is used “*. . . contra llagas sipiliticas’’. Three spe- 
cies may be included from Mexico: Mascagnia septen- 
trionalis (G@. Thurber 901, Malpighiaceae) was used at 
least a century ago, the root being a putative specific 
against syphilis. A ‘‘remedy for gonorrhoea’? comes 
from A eeratum conyzoides (P. C. Standley 19220, Com- 
positae). ‘The macerated leaves of Hintonia latiflora 
G. B. Hinton Herb. No. 3413, 4817, Rubiaceae) are 
taken internally for malaria and pinto. Pinto, or pinta, 
is the common name in Mexico for a form of trepone- 
matosis, an infection closely related to venereal syphilis. 

In Africa, there is Sabicea Vogelu (G. EF. S. Elliot 
4175, Rubiaceae) **‘Used for gonorrhea by Natives. . .”’ 

The third and fourth largest categories of ethnogyn- 
ecological notes pertain to menstruation and ‘‘female 
disorders’’, respectively. ‘These notes deserve special at- 
tention. ‘The plants in question apparently are used to 
treat what appear to be, for the most part, routine func- 
tional and minor ailments. But there is the possibility 
that at least some of these plants act, in fact, as regula- 
tors of reproductivity. As unrecognized birth control 
agents—for instance, as abortifacients or promoters of 
fertility—these species would be of considerable interest 
in connection with human population studies. 

Plants in this category include Pollia thyrsifolia (C. 
rake 491, Commelinaceae) in the Philippine Islands: 
‘*Teaves applied for amenorrhea’’. 

In the West Indies, Vournefortia volubilis GS. Kings 
140, Boraginaceac) is **Used for female trouble’*. Here, 


[ ooQ 
| Oo ‘ 


too, Eleutherine bulbosa (D. Taylor 30, Tridaceae) is 
‘* |. used as acure for irregular menstruation and meno- 
pause’. In Peru, L/. plicata (". Woythowsht 5744) pro- 
duces bulbs from which a decoction is made for treating 
hemorrhages. Bulbostylis capillaris (W. A. Archer 4931, 
Gramineae) from Paraguay is ‘‘Sold by herb dealers in 
market at Asuncién. . .as blood purifier in female dis- 
orders’. In El Salvador, Polypremum procumbens (Dr. 
A. Van Severen s.n., Loganiaceae) is believed to be a 
‘‘Remedy for ‘metritis’ ” 

There are three Mexican Compositae of interest: Hap- 
lopappus spinulosus (Dr. Gregg 6) over a century ago 
was decocted for ‘‘. . . disorders of uterus detention of 
eatamenia’’. Mranseria ambrosioides (H.S. Gentry 1336) 
was used as recently as 1935 for ‘‘female trouble’’, the 
roots being cooked in water. In the same year, it was 
reported of Viguiera montana (H. S. Gentry 1288) that 
‘VV omen put leaves on stomach to cause menses to flow, 
‘por sale la sangre’ ” 

Perhaps of greater interest to students of fertility are 
those few species which are claimed directly to be able 
to control human reproduction. These species represent 
the sixth largest category of ethnogynecological notes. 
They are included at this point, out of sequence, because 
of their relationship with the two foregoing categories. 
One might cite here Vochysia lomatophylla (I. Woyt- 
kowshi 6021, Vochysiaceae), collected in Peru in 1960: 
‘* |. perhaps used by Campa tribe as contraceptive ?”’ 
In the Compositae, Tostephane heterophylla (H. S. Gen- 
try 6349) was reported from Mexico in 1941 to have 
tuberous roots ‘‘... decocted for medicine; makes 
women fertile”’. 

The fifth largest category of ethnogynecological notes 
is made up of plants administered during labor, or in 
childbirth. To here, all examples cited have come from 


[ 339 ] 


genera scarcely known for their utility to man in any con- 
text. ‘Those which follow represent genera which already 
enjoy economic standing of some sort but whose herein 
named uses are not widely known. 

A number of examples is to be found from the Philip- 
pine Islands. An indeterminate species of Loranthus 
(G. EH. Edano 1999, Loranthaceae) bears leaves which 
are pounded ina mortar and applied to the stomach be- 
fore childbirth. Glochidion cauliflorum (M.D. Sulit 3297, 
THuphorbiaceae) is used as follows: ‘‘Decoction of roots 
mixed with roots of cacao and coffee given to women for 
quick delivery’’. An ‘‘... infusion of the roots...”° of 
Ocimum sanctum (G. I. Hdano 1618, Labiatae) ‘*. .. is 
given to mothers for childbirth’’. The use of Ocimum 
reminds one of recent work on Nepeta Cataria’ and Salvia 
divinorum,” both also mints. The last two genera ap- 
pear to contain psychopharmacological constituents. Is 
it possible that Ocimum may produce a similar intoxica- 
tion and be used in primitive childbirth somewhat as the 
plant-derived hypnotic scopolamine has been employed 
in modern labor? 

‘Two species in the Leguminosae might be cited from 
the Caroline Islands: Pterocarpus indicus (C.C. Y. Wong 
419) produces leaves which are pounded to fine particles 


and applied for vaginal ruptures. Women “‘. . . express 
the leaves. ..”” of Phaseolus adenanthus (C.C.¥. Wong 
302) **... and drink the sap for labor pains’’. 


In Ecuador, Helotropium argenteum (I. Prieto 2556, 
Boraginaceae) is employed to prepare an ‘‘Infusion of 
leaves given to women who have fits and spasms during 
childbirth’*. This genus has been recorded previously as 
possessing medicinal properties but not, to my knowl- 
edge, as an antispasmodic.° 

‘The smallest category of ethnogynecological notes re- 
lates to pregnancy. It encompasses only two species, 


[ 840 ] 


from the Gramineae. Pennisetum alopecuroides (C. O. 
Frake 512) from the Philippine Islands has roots which 
are ‘‘.. . applied to stomach to reduce size in pregnancy 
(!)°° Cyperus brevifolius (S. F'. Kajewsht 276) is used in 
the New Hebrides Islands, where it is macerated with 
other plants and drunk for good health during pregnancy. 

I should not leave this subject without a word on 
aphrodisiacs. They have not been included among ethno- 
gynecological notes. Possible aphrodisiacs number possi- 
bly fewer than 15 among all notes from the herbarium 
search. Perhaps the most intriguing entry is little known 
Centropogon calycinus (Y. Mewia 7701, Campanulaceae) : 
in Ecuador, it is ‘‘Used as a love potion’’. Madllotus 
Poilanei (M. Poilane 26084, Euphorbiaceae) belongs to 
a medicinal genus’ but is reported from Indochina as 
having a ‘‘Racine tonique et aphrodisiaque”’. Jatropha 
angusti (J. West 3796, C. Vargas C. 408, Euphorbiaceae), 
also from a medicinal genus,’ is reputed to be an aphro- 
disiac in Peru. Ptychopetalum olacoides (G. Stahel 269, 
Olacaceae) is supposed in Surinam to act as an aphro- 
disiac; the species also is the source of Muira puama, 
employed in medicine.’ 

Among alleged love charms are species of the genera 
Loranthus (Loranthaceae), Aristolochia (Aristolochia- 
ceae), Canavalia (Leguminosae), and Premna (Verben- 
aceac). Sundry common names suggest aphrodisiacal 
properties: among these I have included plants called 
‘Tove Bush’’, “Amor Seco’’ and ‘‘Matrimonio Viejo”’. 

There is one antaphrodisiac: of Davilla lacunosa (LO. 
Williams 5098a, Dilleniaceae) in Brazil, it is said: ‘Give 
tea to animals and they are impotent’’. 

In conclusion, I want to say that species of plants 
associated with ethnogynecology warrant careful inspec- 
tion by modern gynecological and obstetrical pharma- 
cologists. I would emphasize further that, in considering 


[ B41 J 


these species, one must also expand one’s view of their 
possible usefulness beyond the purposes for which they 
are employed in the simplistic contexts of their ethno- 
botanical origins. ‘Today we can cure venereal disease ; 
we have anodynes for dysmenorrhea and synthetic opi- 
ates for labor. The major value of the notes referred to 
may lie in how they might bear incidentally upon such 
fields as cancer, heart, mental and metabolic research. 
It is in such a questioning posture that we hope to present 
these and all our materials to science in a forthcoming 
catalogue. 

Today, not even ten years after our first thoughts on 
searching herbaria, we inevitably ask whether it is not 
imperative to attempt now—perhaps through the inter- 
national cooperation of various scientists and govern- 
ments—to preserve living examples of all extant species, 
including economic and presently non-economic plants. 
| take this opportunity to recommend that coordinated 
efforts to this end would be particularly fruitful in the 
so-called underdeveloped nations where, it so happens, 
one encounters the most promising of ethnobotanical 
raw materials, directly from the indigenous societies 
which give birth to their uses. If civilized man does not 
act now, he may leave his descendants irretrievable losses 
and, possibly, the undoing of his own species through 
want of some missing botanical essential to a future 
generation. 


[ 342 ] 


Gr 


LITERATURE CITED 


. von Reis, S. 1962. Herbaria: sources of medicinal folklore. Econ. 


Bot. 16: 283-287. 


Altschul, S. von Reis. 1967. Psychopharmacological notes in the 
Harvard University herbaria. Lloydia 30: 192-196. 


Altschul, S. von Reis. 1968. Unusual food plants in herbarium 
records. Econ. Bot. 22: 293-296. 


Altschul, S. von Reis. Ethnopediatric notes in the Harvard Uni- 
versity herbaria. ined. 


Uphof, J. C. Th. 1959. Dictionary of economic plants. Weinheim, 
Germany: J. Cramer. 


Crosby, A. W. 1969. The early history of syphilis: A reappraisal. 
Amer. Anth. 71: 218-227. 


Waller, G. R., G. H. Price and FE. D. Mitchell, 1969. Feline at- 
tractant, cis, trans-nepetalactone: metabolism in the domestic cat. 


Science 164: 1281-1282. 


Epling, C. and C.D. Jativa-M. 1962. A new species of Salvia from 
Mexico. Bot. Mus. Leafl. Harvard Univ. 20: 75-76. 


Wasson, R.G. 1962. A new Mexican psychotropic drug from the 
mint family. Bot. Mus. Leafl. Harvard Univ. 20: 77-84. 


[ 343 ] 


DE PLANTIS TOXICARIIS E MUNDO 
NOVO TROPICALE COMMENTATIONES VII 
SEVERAL ETHNOTOXICOLOGICAL NOTES 
FROM THE COLOMBIAN AMAZON 
BY 
Ricnarp Evans SCHULTES 


Many of our ethnotoxicological observations made 
amongst Indians of the northwest Amazon indicate in- 
teresting uses of plants the genera of which have never 
been chemically investigated. The several species dis- 
cussed below fall very definitely into this category and 
would seem to represent areas profitable for phytochemi- 
‘al enquiry. 

These notes are a continuation of investigations into 
poisonous plants of the New World tropics which I have 
been carrying out since 1941 and which is currently being 
supported by a grant from the National Institutes of 
Health (No. I.M-GM00071-01). 

The voucher specimens cited are preserved in the Gray 
Herbarium, the Arnold Arboretum and the HKconomic 
Herbarium of Oakes Ames of Harvard University or in 
the Herbario Nacional Colombiano. The illustrations 
were prepared by Mr. Josua B. Clark and the late Mr. 
John Stanwell-Fletcher. 

ARACEAE 

No chemical studies appear to have been carried out 

on the genus Urospatha. The juice from the rhizome of 


[ 345 | 


t Brazilian species—Uvrospatha caudata Schott—is used 
medicinally in the treatment of skin troubles (Peckholt, 
T.: Pharm. Rundshau 10 (1892) 279, 11 (1893) 879). 


Urospatha sagittaefolia Schott Aroid. (1853) 4. 


Cotombia: Comisaria del Amazonas, Rio Amazonas, Leticia and 
vicinity. “‘In swamp. Spathe green externally, white-green within, 
Stem mottled brown-green. Height 6 feet.’> August 29-September 
12, 1966. RE. Schultes, R.F. Raffauf et D. Soejarto 24039. 


A field spot test with modified Dragendorf reagent in- 
dicated that Urospatha sagittaefolia is alkaloid-positive. 


Urospatha somnolenta PF. EL. Schultes in Bot. 
Mus. Leafl. Harvard Univ. 18 (1958) 123. 


Cotompra: Comisaria del Vaupés, Rio Kuduyari, Cachivera de Ita- 
pinema. August 14, 1960. R.E. Schultes 22589, 


This is the second collection referable to Urospatha 
somnolenta. Like the plant from the type locality, 
Schultes 22589 grew alongside a quartzitic rapids in 
swampy holes in the rock. The locality of this second 
collection belongs geologically to the same formation as 
the type locality. 

The Kubeo Indians at Yapoboda consider the root of 
Urospatha somnolenta to be toxie if ingested, yet the 
ashes of the entire plant are employed medicinally in the 
form ofa poultice on ulcers and intected cuts. The Kubeo 
name of the plant is 6-me’-na. 


MyRrsInackEAE 
Conomorpha citrifolia J/ez in Engler Pflanzenr. 
4 Myrsin. (1902) 256. 


CotomBia: Comisaria del Vaupés, Rio Paraparand, Raudal Na-hoo- 
¥ ee on 8 . aes . 
gaw-he. Flowers whitish yellow. Small tree along bank.*’ Septem- 
ber 1952. R.E. Schultes & 1. Cabrera 17593. 


[ 346 ] 


The Puinave Indians, some of whom have migrated 
into the Apaporis basin, call this treelet yoom-dd-ha and 
rasp the bark into chicha to impart to it a rather peppery 
taste. It is recommended also as a febrifugal tea. 


APOCYNACEAE 


Ambelania Lopezii JMoo0dson ev RoE. Schult. in 
Bot. Mus. Leafl. Harvard Univ. 15 (1951) 76. 


Cotomsi1a: Comisaria del Vaupés, Rio Guainia, below San Antonio, 
Cafio del Loro. ‘*Molongé. Flowers white, fragrant. Growing at edge 
of water.’’ June 1948. R.E. Schultes et F. Lopez 10152,—Comisaria 
del Vaupés, Rio Kananari, Cachivera Palito. “"Bush. Flowers fragrant, 
white.”’’ July 25,1951. RE. Schultes et I. Cabrera 13175.—Rio Apa- 
poris, Raudal Yayacopi (La Playa) and vicinity. “‘Small tree. Flow- 
ers white, very fragrant. Latex white.’” August 18, 1952. Schultes 
et Cabrera 16968.—Rio Piraparand, middle course. **Four meters tall. 
Flowers white, fragrant.’’ September 1, 1952. Schu/tes et Cabrera 
17135,—Rio Piraparand, Cano Teemeefia. *‘Large bush. Flowers 
white, fragrant. Latex white.*’ September 6, 1952. Schultes et Cabrera 
17245. 


The Barasana Indians boil the leaves of this bushy 
treelet with the bark of Destictella racemosa and Mar- 
tinclla obovata to make one of their arrow poisons. 

Amongst the related Makunas of the same area, who 
know this plant as wey-gazw-76-mee’-h6, it is apparently not 
employed for this purpose but reputedly as a fish poison. 


BIGNONIACEAE 


Distictella racemosa (Bur. & K.Schum. ex Mart.) 
Urban in Kedde Repert. 14 (1916) 810. 

CotomBia: Comisaria del Amazonas, Rio Apaporis, Soratema (above 
mouth of Rio Kananari) and vicinity. Alt. about 900 feet. ““Liana. 
Flowers cream-white. July $31, 1951. R. BE. Schultes et I. Cabrera 
13215,.—Same locality. ““Woody vine. Flowers white. Barasana name 


= mee-tsee-boo-koo’-na.** January 28, 1952. R.E. Schultes et I. Cabrera 
14990. 


The bark of Deistictella racemosa is one of the ingredi- 


[ 347 ] 


PLaTE LXXX 


vf) 


LU TNS 


AMBELANIA  Lopezii Woods 


JBC 


[ 348 | 


Puate LXXXI 


i Mh 


ig F 
; e Bis 


Ambelania Lopezii Woodson in a flooded caatinga near the conflu- 
ence of the Rios Guainia and Negro, Colombia. 
Photograph: R.E. Scuuites 


PLATE LXXXII 


DISTICTELLA racemosa ? 
(Bur.et Ar 


ents of atype of curare prepared by the Barasana Indians 
living on the Rio Apaporis near the mouth of the Kan- 
anari. ‘he other plants said to be used in this preparation 
are the leaves of -Ambelania Lopez and the bark of 
Martinella obovata. The root of Distictella racemosa is 
reputedly very toxic. 


Martinella obovata (H BA.) Bureau et BK. Schu- 
mann ev Martius F). Bras. 8, pt. 2 (1897) 161. 
CotomsBia: Comisaria del Amazonas, Rio Apaporis, Soratama (near 


mouth of Rio Kananari), January 28, 1952. R.E. Schultes et I, Cabrera 
14994. 


The bark of Martinella obovata, together with the 
leaves of Ambelania Lopezu and the bark of Distictella 
racemosa, is utilized by the Barasana [ndians in elaborat- 
ing on arrow poison. An infusion of the bark is said to 
be a febrifuge but dangerous to use. 


[ 351 ] 


PLatTE LXXXIII 


MARTINELLA obovata (HBK) Bur. & K. Schum. 


JBC 


[ 352 |